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Printed in Great Britain at the University Press ^ Cambridge 
{Brooke Crutchley^ University Printer) 
and published by the Cambridge University Press 
{Cambridge^ and Bentley House, London) 

Agents for US, A,: The University of Chicago Press 
Canada and India: Macmillan 

The British 
Mycological Society 

{Recognosce notum, ignotum inspice) 


Volume XXXI 

Edited by 

B. Barnes and W. C. Moore 



Vol. XXXI, Parts I & II 

July 1947 



Department of Botany^ Central College^ Bangalore^ India 


(With 9 Text-figures) 

Species of the genus Sphaceloma de Bary, with their ascigerous stage 
I Elsinoe Racib . , have received a good deal of attention from plant pathologists 

! on account of the scab or anthracnose disease they produce on plants of 
? economic importance. Recent monographic studies by Dr A. A. Bitan- 
[ court and Dr A. E. Jenkins have greatly advanced our knowledge of the 
» group. In India, only two Sphaceloma ^tdigcs are known: one is S. Fawcettii 
^ Jenkins, which is the conidial stage o^ Elsinoe Fawcettii Bitan. & Jenkins, 
f recorded on species oi Citrus (Jenkins & Fawcett, 1933) and on Hesperethusa 
crenulata^ from Bengal (Jenkins, 1936) ; the other is the Sphaceloma stage of 
Elsinoe Bitancourtiana Thirumalachar, recently described by me (1946). In 
collections made in Mysore, four more species of Sphaceloma were found, 
some of the hosts being plants of economic importance. Further studies are 
being made to investigate the possible existence of the ascigerous stages. 

; An account of the Sphaceloma stages is presented here. 

(i) A Sphaceloma msEASE oe Santalvm albvmIj. 

Santalum album, the s andal tree, is of great economic importance on account 
of the valuable essential oil contained in the heart wood. Diseases of the 
sandal tree have been the subject of several detailed investigations in India. 
A species of Sphaceloma was found parasitizing the leaves in some of the 
collections made by me in the Mysore forests. The infected leaves have a 
warty surface with copper-coloured patches on the upper surface, and tend 
to roll and curl inwards. In association with Asterina congesta Cke. and an 
Oidium, which are also found as leaf parasites on Santalum album, the damage 
caused by the Sphaceloma is important, for the affected leaves fall long before 
healthy leaves. As no detailed search has been made in the sandal-growing 
areas of Mysore for studying the spread of the disease, it is not possible to 
evaluate the extent of the damage. 

The infection first appears as cinereous areas on the upper surface, which 
gradually turn purplish to black, appearing as discolorations. The acervuli 
are intra-epidermal in origin and are distributed on either surface of the 
leaves. They rarely coalesce, and even where they are in close proximity 
the demarcation between the individual sori is still visible. The conidio- 
phores are developed from scanty stromata or hyphal cells which become 
indistinguishable in later stages. The conidiophores are grouped in plumose 
tufts (Fig. i), 43-60 ju, long and 2-2 broad, hyaline, thin-walled and 
pointed at the apex. The conidia are produced in succession at the tip. 
They are hyaline, thin-walled, ovate to ellipsoid (Fig. 2), measuring 

YoL XXIX, Part IV: issued 21 January 


2 Transactions British Mycological Society 

5-9 X 2-5-3JU.. No sptcits of Sphaceloma has so far been described on any 
of the members of the Santalaceae, and comparison with the described 
species indicates that the species on Santalum is new. 

Sphaceloma Santali Thirumalachar, n.sp. 

Producing cinereous patches on the upper leaf surface, sometimes with 
purplish to black tinge and appearing as brown to black discolorations. 
Acervuli amphigenous, but mostly epiphyllous, intra-epidermal, 60-140/x 
broad, ^S-^Op. high, not coalescing with one another; conidiophores 
43-60 p. long, produced in plumose tufts and developed from scanty 
stromata or hyphal cells which are indistinguishable later on; conidia 
produced acrogenously, hyaline, ovate-elliptic, measuring 5-9 x 2-5-3 p. 

Habitat. On the leaves of Santalum album L., Lakkavalli, Mysore State, 
leg. M. J. Thirumalachar, 14 October 1945. Type deposited in the Herb. 
Crypt. Ind. Orient., New Delhi, in the Imperial Mycological Institute, 
Kew, England, and in the Institute Biolo^co, Sao Paulo, jlrazil. 

Producit cinereas maculas in supeifore facie foliorum, nonnumquam 
palhde purpurescentes vel nigras, quae et brunneo-nigrae discolorationes 
apparent. Acervuli intra-epidermales, amphigeni sed ut plurimum 
epiphylli, latitudine 60-140P, altitudine 43-60 p, baud coalescentes inter 
se; conidiophori 43-60P longi, plumose caespitosi, evoluti ex stromatis \-el 
hypharum exiguis cellulis quae postea indistinctae evadunt, 43-60P longi, 
apice acuto; conidia acrogene producti in apice, hyalina, ovata vel elliptica, 
magnitudinis 5-9 X 2-5-3 p. 

In folds Santali albi L., Lakkavalli, Mysore State, leg. M. J. Thiru- 
malachar, 14. ro. 1945. 

(2) A NEW Sphaceloma on Osyris Wightiana Wall. 

Osyris Wightiana, a member of the Santalaceae, is a partial root parasite in 
the hilly districts of Mysore. The heart wood of the plant is very faintly 
fra^ant, and is often used for adulterating sandal wood. Some collections 
of the leaves of this plant made near Nandi Hills, Mysore State, posses.scd 
co^cent^c leaf spots in association with the epiphyllous ascomycetoiis 
hm^m Mehola Osyndicola Hansf. which forms large sooty blotches masking 
the irfection spots of the Sphaceloma. The infection spots are minute^ 
amphigenous, arranged in perfect concentric rings (Fig. 3). The oldest 
acervuli are situated in the centre, the infection spreading centrifugally. 
of Sheaves ^ appear as minute depressions on the su rface 

spot show that the .sori are intra- 
epidermal, formed by the Rouping of a stroma of hyphal strands (Fig. .t . 

turn beneath the son become depleted of their contents and 

turn orange coloured and ca^er-hke. Palisade layers of conidiophores 
hat are differentiated froip the stroma, break through the upper wall of 
the epidermis and the cuticle, and abstrict off conidia. The maUrc conidi-i 
are hyahne, one-celled, ovate to globose (Fig. 5) and measure 1 x - - I 

the Mature oAL: ri 

and other characters, such as the length of the com'diophores 

Some new Sphaceldma diseases of economic plants in Mysore 3 

Sphaceloma Osyridis Thirumalachar, n.sp. 

Infection spots amphigenous, appearing as minute depressions, arranged 
in perfect concentric rings. Acervuli intra-epidermal, composed of strands 
of hyphae and a palisade layer of conidiophores, not coalescing with each 
other and not becoming continuous. Conidia hyaline, acrogenous, ovate to 
spherical, thin-walled, measuring 3 x 2*5/x. 

Habitat, On the leaves of Osyris Wightiana Wall., Nandi Hills, Mysore 
State, leg. M. J. Thirumalachar, 19 February 1944. Type deposited in the 
Herb. Crypt. Ind. Orient., New Delhi, in the Imperial Mycological 
Institute, Kew, England, and in the Institute Biologico, Sao Paulo, Brazil. 

^ Infectionis maculae amphigenae, apparentes ut minutae depressiones, 
dispositae in circulis perfecte concentricis, Acervulus intra-epidermalis, 
constans ex hypharum fasciculis atque strato conidiophororum, non 
coalescens cum aliis nec cum illis continuus evadens. Conidia hyalina, 
acrogena, ovata vel sphaerica, tenuiter parietata, magnitudinis 3 x 2 '5//-. 

In iolm Osyridis Wightiana in loco Nandi Hills, Mysore State, leg. 
M. J. Thirumalachar, 19. 2. 1944. 

(3) An anthracnose disease of oleander leaves 

Collections of oleander leaves made by me near KoppaRoad, Mysore, were 
severely parasitized by a Sphaceloma, Another collection of the same fungus 
made near Kemmangundi, a hill station in Mysore, was also examined. 
The lesions on the leaves were circular to irregular, whitish, bordered by 
a brownish black margin (Fig. 6) . By the coalescence of these lesions, the 
infection patch enlarges and occupies most of the leaf surface. When 
examined with a lens, the surface of the infected patch presents a punctiform 
to verrucose appearance. In severe infection, there is heavy defoliation of 
the plants. 

The acervuli are amphigenous but mostly epiphyllous and are grouped 
on the infection spots. They are intra-epidermal, formed by the grouping 
of the hyphal strands within the epidermis. As is well known, the epidermis 
in Nerium oleander possesses two to three layers and it has been observed that 
the acervuli are formed in all three layers. Especially when the sori are 
formed within the innermost layer of epidermis, the developing sorus breaks 
through the two superposed epidermal layers, appearing as subepidermal 
in origin. The acervuli coalesce when they are developed in close proximity 
and thus become continuous. 

The fundaments of the sori are formed by the grouping of the hyphae 
which form a thick well-developed stroma of several tiers of cells (Fig. 7). 
The conidiophores are indistinguishable from the rest of the cells of the 
stroma, except that they are the terminal group of cells. These rupture the 
upper wall of the epidermis and develop acrogenously hyaline, spherical 
conidia (Fig. 8) which are thin-walled and measure 4-5 x 2 •5-4 ft. No 
species of Sphaceloma has previously been reported to occur (m Mefum or its 
allied genera and the present species cannot be accommodated in any of the 
previous species so far described. 

Some new Sphaceloma diseases of economic plants in Mysore 5 

Sphaceloma Oleander! Thirumalachar, n.sp. 

Leaf spots spherical to irregular, densely grouped, 1-4 mm. in diam. 
distributed over the entire surface of the leaves, whitish with a brownish black 
margin, slightly elevated and appearing rugged owing to the punctiform 
masses of Sphaceloma. Acervuli are intra-epidermal, formed in the first, 
second or the third layers of the upper epidermis, seldom amphigenous, often 
becoming coalescent, 140-200 /x in diam. and 60-85 /x high; stroma well 
developed with several tiers of cells; conidiophores short, 5”6‘5 x 2-2;5^, 
breaking through the upper wall, developing acrogenously hyaline, thin- 
walled conidia which measure 4-5 x 2*5-4 /x. 

On the leaves of Nerium Oleander ^ Koppa Road, leg. M. J. Thirumalachar, 
3 April 1945, leg. H. G. Govindu, Kemmangundi 10 October 1945 (Type). 
Type deposited in the Herb. Crypt. Ind. Orient., New Delhi, in the 
Imperial Mycological Institute, Kew, England, and in the Institute 
Biologico, Sao Paulo, Brazil. 

Foliorum maculae sphaericae vel irregulares, dense aggregatae, 1-4 mm, 
diam., dispersae per totam foliorum faciem, albidae margine fusco-nigro, 
tenuiter elevatae atque asperae ob punctiformes massas Sphacelomatis. 
Acervuli intra-epidermales, producti intra primum, secundum vel tertium 
stratum epidermatis superioris, raro amphigeni, saepe inter se confluentes, 
ut plurimum 60-85 x 140-200 /x, stromate bene evoluto multiplice 
cellularum ordine; conidiophori breves, 5-6*5 x 2-2*5 /x. perforantes 
exteriorem epidermatis parietem; conidia acrogene producta, hyalina, 
tenuiter-parietata, magnitudinis 4-5 X 2*5-4/x. 

In foliis Merit Ofean&n; Koppa Road, Mysore, leg. M. J. Thiru- 
malachar, 3. 4. 1945; leg. H. C. Govindu, Kemmangundi, 10. 10. 1945 


Indigenous species of Curcuma are well distributed in the forests of Mysore, 
The rhizomes of many of these wild turmerics are used as an additional 
source of starch by the local people. In one species of Curcuma it was 
observed that the leaves had developed numerous small lesion spots, which 
on examination proved to be caused by an undescribed species of Sphace- 
loma. The lesions are brownish yellow, circular to ovate, 2-4 mm. in diam. 
and mostly found on the upper surface. 

Sections through these infection spots revealed that the acervuli aref 
subcuticular (Fig. 9) and very rarely, and then probably exceptional. 

Legends for Figures 1-9 

Fig, I. Ac^xviAns or Sphacdoma SantalL (X500,) 

Fig. 2. oi Sphaceloma Santalu (xiooo.) 

Fig. 3. Showing the ixdtciion ot Sphaceloma Osyridis. (Nat. size.) 

Fig. 4. Actxynlm oi Sphaceloma O^ridis. ( X500.) 

Fig. 5, Si^Qxts or Sphaceloma Osyridis, { x 1^00.) 

Fig. 6. Anthracnosed leaf of Wmum (Nat. size.) 

Fig. 7. Showing the intra-epidermal acervulus in Sphaceloma Okanderi. ( x 250 ) 
Fig. 8, Spores, (x iooo.) 

Fig. 9. Acorvnlm of Sphaceloma curcumae. ^00.) 

6 Transactions British Mycological Society 

intra-epidermal. A compact layer of somewhat pale pseudopareii- 
chymatous cells is produced. From this basal stroma, palisade layers of 
conidiophores are differentiated which are fusiform and become pointed at 
the apex. The conidiophores become laterally united and thus present a 
compact appearance. Most of the acervuli were found to be immature 
and only very young spores in the process of abstriction at the tip have 
been observed. Consequently the size and shape of the mature conidia 
remain unknown. The host cells and the entire stroma of the acervulus 
contained numerous microconidia. Such microconidia were found by 
Jenkins and Bitancourt in a number of species oiElsinoe. This is apparently 
the first record of Sphaceloma on tht Zingiberaceae. 

Sphaceloma Curcumae Thirumalachar, n.sp. 

Acervuli subcuticular, epiphyllous, round or irregular, minute, 31-50/x 
in diam., pale yellow, composed of compact pseudoparenchymatous cells 
with conidiophores which are palisade-like and coalesce laterally, 20-25/i 
long; conidia not observed, microconidia numerous and filling up the cells. 

On leaves of Curcuma sp., Kemmangundi, Mysore State, leg, M, J, Thiru- 
malachar, 9 October 1945. Type deposited in the Herb. Crypt. Ind. 
Orient., New Delhi, Imperial Mycological Institute, Kew, England, and in 
the Institute Biologico, Sao Paulo, Brazil. 

Acervuli subcuticulares, epiphylli, globosi vel irregulares, minuti, 3 i~50|x 
lati, alti, pallide lutei, compositi ex compactis pseudoparenchyma- 
ticis cellulis, conidiophoris vallatis atque lateraliter coalescentibus, 20-251X 
longis ; conidia haud visa, microconidia plurima, cellulas implentia. 

In foliis Curcumae sp., Kemmangundi, Mysore State, leg. M, J, TMru- 
malachar, 9, 10. 1945. 


Jenkins, A. E. (1936). A Sphaceloma on fruit Hesperethusa crmulata, a remott^ Ciirus 
relative from India, Fhytopathologp^', 

Jenkins, A. E. & Fawcett, H. S. ( ^933). Records ofeitrus scab from herbarium specimens 
of the genus Citrus in England and the United States, Phytopathology^ xxiii, 475-82. 
Thirumalaghar, M. j. (1946). An undescribed species of Elsinoe from Mysore. i/>w- 
xxxviii, 220-5. 

{Accepted for publication 10 October i 

[ 7 ] 



Department of Botany], Central College^ Bangalore^ India 

(With 8 Text-figures) 

The fungi associated with or parasitizing the thalli of the liverworts have 
attracted the attention of numerous investigators. A brief review of the 
various facts pertaining to this subject is given by Nicolas (1932) . The fungi 
are usually found only in association with the gametophytes and it is 
uncertain if they are weak parasites or symbionts. The possible occurrence 
of a my corrhizal fungus, within the thallus of Marchantia was pointed out 
by Kashyap (1916) in India^ and recently Mahabale & Bhate (1945) have 
recorded the presence of a mycorrhizal fungus in the ventral tissue of the 
liverwort Fimbriaria angusta St. As regards the disease-producing fungi on 
liverworts in India, the life cycle of Phaeospkaerella Ricciae F. L. Steph. 
parasitizing the gametophytes of Riccia himalayensis St. was described by 
Srinivasan (1939). In the course of the collection of material of Hepa- 
ticeae by me for class work, the occasional presence of fungi as parasites 
was noticed. Since no fungi were suspected to be present at the time of 
collection, the materials were preserved in formalin alcohol and conse- 
quently they were unfit for undertaking any germination or cultural 

(i) A Cladocbytrium parasitic on Aneura sp. 

In some of the collections of gametophytes of sp. made near 

Agumbe, Mysore State, many of the thalli had developed discoloured 
patches. In place of the healthy green colour, the affected patches had 
turned straw yellow. A detailed examination of such thalli indicated that 
they were parasitized by a species of Cladochytrium, lAo species of Clado-- 
chytrium has so far been recorded from India, but another genus of the 
family Nowakowskiella ramosa Butler, was reported on 

decaying wheat stems by Butler (1907). 

The fungus is intramatrical, the ‘rhdzomycelium’ pervading the cells of 
the host and forming extensive branches. At short intervals the rhizo- 
mycelium swells up and forms spindle-shaped swellings or the turbinate 
organs (Figs, i, 2) which are very characteristic of the family Glado- 
chytriaceae. These spindle-organs are mostly continuous (Fig. 2) without 
any wall formation, but sometimes (Fig. i) they are definitely septate. It 
might therefore be well to consider the spindle organs in the present species 
as being sparsely septate. In distinguishing between the genera 
chytrium and HowakowskieUa^ th.Q presence of an operculum in the zoo- 
sporangium of the latter is stated to be an important character, and Whiffen 
(1943) considers that the presence of the septations in the spindle-organs 
of Cladochytrium 2idds another differentiating character. The spindle-organs 
of are therefore regarded as unseptate and continuous. 

8 Transactions British Mycological Society 

Zoosporangia and resting spores were observed in large numbers. Tlie 
zoosporangia are terminal or intercalary, their size and shape varying 
according to the host cell in which they are lodged. The division of the 
contents of the zoosporangium into zoospores was frequently noticed 
(Fig. 3)5 but their actual discharge could not be followed on account of the 
preserved state of the material. On some old zoosporangia, small papil- 
late structures which might represent the remnants of the exit tubes were 
seen. The resting spores are ovate to spherical, cinnamon-yellow, thick- 
walled and smooth. The mature spores are often formed by the trans- 
formation of the spiiidle-organs and measure 30-44 /x in diam. 

While species of Cladochytrium^ such as C. replicatum Karling, are considered 
to be very weak parasites or almost saprophytes, other species, such as 
C. Mowakowskii Sparrow (Sparrow, 1931), is stated to be a virulent 
parasite on algae. The discoloration produced in the thalli of Aneura by the 
species under study shows that it might also be a weak parasite. Inocu- 
lation experiments carried out by Karling (1931) indicated that the species 
of Cladochytrium wide host ranges and the identity of the species 

therefore on the basis of host alone would be erroneous. C. replicatum^ for 
instance, has been successfully inoculated into several hosts, including the 
livtvwovt Anthoceros (Karling, 1931)- That being so, the identity of the 
species should be based on morphological characters and spore measure- 
ments alone. The sparsely septate nature of the spindle-organs, the 
inconspicuous, papillate exit-tube and the difference in size of the resting 
spores as compared with those of most of the species of Cladochytrium^ has 
led me to the view that the species may be an undescribed one, though 
future investigations may reveal it to be identical with some well-established 

Cladochytrium Aneurae Thirumalachar n.sp. 

Thallus intramatrical consisting of fine tenuous branched rhizo- 
mycelium with numerous spindle-shaped swellings which rarely become 
septate; zoosporangia spherical to oval, depending upon the shape the 
host cell, often with a single constricted exit pore. Resting spores terminal 
or intercalary, often formed by the transformation of spindle-organs, 
usually intramatrical, ovoid to spherical, cinnamon-yellow, thick-walled, 
smooth, measuring 30-44 /X in diam. 

Habitat, As a weak parasite within the thallus of Amura sp. (Jimger- 
manniales), Agumbe, leg. M. J, Thirumalachar, 4 April 1945. Type de- 
posited in the Herb. Crypt. Ind. Orient., New Delhi, and in the Imperial 
Mycological Institute, Kew, England. 

Thallus intramatricalis constans ex gracili tenuique atque ramose 
rhizomycelio ornato^ plurimis fusiformibus tumoribus qui rare septati 
evadunt; zoosporangia sphaerica vel ovata pro forma cellulae parasitatae, 
52— 100 /X diam., unico constricto exitus poro. Quiescentes sporac termi- 
nales vel intercalares, saepe evolutae ex transformatione fusiformium 
organorum, plerumque intramatricales, ovatae vel sphaericae, cinna- 
momo-luteae, crasso pariete ornatae, leves, magnitudis 30-44 /x. 

Fungal diseases of hryophytes in Mysore. M.J. Thirumalachar 9 

Parasitus debilis intra thallum Aneurae sp. (Jungermanniales) in loco 
Agumbe; leg, M. J. Thirumalachar, 4. 4, 1945. 

(2) Olpidiopsis Ricciae Du Plessis 

Olpidiopsis Ricciae Du Plessis was first described by Du Plessis (1933) within, 
the rhizoids of jSzrrfa in South Africa. Large numbers of sporangia and 
oospores were observed by him within the normal and pegged rhizoids. 
The presence of a companion-cell which in fact represents the empty 
antheridium attached to the oospore is a characteristic feature. The 
oospores have been observed within the rhizoids and rarely within the 
epidermal cells of Riccia^ and the host range of the fungus is as yet unknown. 

An Olpidiopsis which seemed to be identical with 0 . Ricciae Du Plessis 
was found by me within the rhizoids of many Hepaticeae in addition to 
Riccia. The oospores have been found in the rhizoids of species of 
Marckantiay Plagiochasma^ Fimbriariay Aneura and Notoihylas, It seems 
therefore probable that the fungus may have a wide host range. 

Most of the oospores found were mature. They are yellowish brown with 
a sinuous outer wall. The companion-cell is mostly lacking from the mature 
spores; Du Plessis also records this. Whiffen (1942) states that Olpidiopsis 
Aphanomycis Cornu lacks the characteristic companion-cell. When the 
oospores are present in large numbers it seems that the cell contents of the 
rhizoids become depleted and shrivel up. Sometimes, as in the rhizoids of 
Aneura^ at the place where the young oospore is lodged, the rhizoid turns 
and twists, forming a nodular structure indicating a definite necrotic effect 
(Fig. 5). In other specimens the definite shrivelling up of the rhizoid from 
the free end is noticeable. It might be of interest to take into account the 
view put forward by Du Plessis about the relationship. Because there were 
no visible signs of necrotic effect produced by the oospores, Du Plessis 
suggested that the relationship between the host and Olpidiopsis Ricciae 
might be one of symbiosis. My observations on what seems to be the same 
species of Olpidiopsis indicate that there is weak parasitism as in Clado- 
chytrium^ becoming discernible in certain hosts or when the host is not able 
to offer resistance. 

In studying the host ranges for the species of Olpidiopsis^ both Shanor 
(1940) and Whiffen (1942) pointed out that it is impossible to transfer any 
species of Olpidiopsis from the species of host on whiclx it was collected to the 
host of any other species. No such studies have been made in the present 
investigation to confirm this interesting observation, but, because of the 
close morphological resemblance of the oospores found in the rhizoids of 
the various liverworts, the fungus in all the hosts is referred to 0 . Ricciae. 

(3) A Leptosphaeria parasitiq on Porella 

Species of Porella are very commonly found as epiphytes on the branches 
and leaves of forest trees. In some of the collections of made in 

Nandi Hills, Mysore State, the leafy lobes of the gametophyte were 
noticed to have been parasitized by an ascomycetous parasite, the infected 
leaves wilting away and finally drying up. Examination of the infected 

Transactions British Mycologtcal Society 

« 9 «. \J 


Figures 1--8 

. Showing intramatrical thallus of Ciadockpirium Amurae. ( x 400,) 

. Some of the unseptate spindle-organs. ( X 600,) 

. Stages in the fomiation of zoosjporangium. ( x 800.) 

. Oospores of OIpMops^ Rkcm within ventral tissue Amura. ( x looo.) 
. Oospores of Olpidiopsis Rkciae within the rhizoids, ( x 500.) 

. Gametophyte of Porella with leptosphaerm infection spots, ( x 
, A&om of Leptosphaerm Porellae. ( x8oo,) 

. Ascospore. (xi8oo.) 

Fungal diseases of hryophytes in Mysore. M.J. Thirumalachar ii 

leaves with a field lens, reveals numerous black specks distributed on the 
dorsal and ventral lobes as well as the amphigastria of the thallus (Fig. 6). 
The infection has been traced to a species of Leptosphaeria. The mycelium 
is intercellular, slightly dark, closely septate and ramifies within the tissues 
of the mesophylL Numerous perithecia are distributed on the surface of 
the infected portions. They are minute, black, immersed within the thallus 
with the ostioles projecting as minute black specks. The asci are cylindrical, 
obovate at the apex, and associated with paraphyses. The ascospores are 
fusiform, rounded at both ends, hyaline, and usually three-septate. The 
spores are arranged in a biseriate manner and measure 15-20 x 2*5- 
3*5//, (Figs. 7, 8). The association of the Phoma stage of the fungus with 
hyaline spores was observed. As no species of Leptosphaeria is so far known 
on any of the species of Porella or any other liverworts, it is not possible to 
institute any comparison. The name Leptosphaeria Porellae is proposed for 
the fungus. 

Leptosphaeria Porellae Thirumalachar, n.sp. 

Perithecia on the foliose portions of the thallus, amphigenous, immersed 
in the early stages, later on becoming erumpent; black, spherical, ostiolate, 
ostiole projecting as a small papilla. Asci cylindric, 8-spored, 57-66 x 6*5- 
7*5//,, paraphysate; paraphyses numerous, simple, filiform; spores one to 
two-seriate, pale yellow, fusoid, rounded at both ends, smooth, slightly or 
not constricted at the septa, measuring 15-20 x 2-5-3*5//,. 

Habitat. On the thallus and causing the wilting (^Porella sp., Nandi Hills, 
Mysore State, leg, M. J. Thirumalachar, loFebruary 1944. Type deposited 
in the Herb. Crypt. Ind. Orient., New Delhi, and in the Imperial Myco- 
logical Institute, Kew, England. 

Perithecia super foliosas partes thalli, amphigena, initio immersa, postea 
erumpentia, nigra, sphaerica, ostiolata, ostiolo eminente ut papilla parva. 
Asci cylindrici, 8-spori, 57-66 x 6*5-7*5/x, paraphysati; paraphyses plures, 
simplices, filiformes ; sporae semel vel bis seriatae, pallide luteae, fusiformes, 
utroque apice rotundatae, leves, tenuiter vel nullo modo constrictae ad 
septa, magnitudinis 15-20 x 2*5-3*5/x. 

In thallo Porellae spec., quern marcescentem reddit; in loco Nandi Hills, 
Mysore State, leg. M. J. Thirumalachar, 10, 2. 1944. 

In conclusion the writer wishes to express his deep debt of gratitude to 
Rev, Father H. Santapau, Professor of Botany, St Xavier’s College, 
Bombay, for kindly rendering into Latin the diagnosis of the new species. 
Grateful thanks are due to Dr B. B. Mundkur and Dr L. N. Rao for kind 
encouragement and valuable suggestions. 


Butler, E. J. (1907). An account of the gtnm Pythitm and some Chytridiaceae. Mem. 
Dep. Agric. India^t^ 

Karling, J. S, (1931). Studies in the Chytridiales. VI. The occurrence and life history 
of a new species of Cladochytrium in cells of Eriocaulon septangulare. Amer. J. Bot. 

. 'XVIII, 526-57.' '■ 

Kashvap, S. (1916). J . Bombay Nat. Hist. Soc. xxiv, 349. 

12 Transactions British Mycological Society 

Mahabale, T. S, & BHATEy P, D. (1945). The structure and life history of Fimbriaria 
dngusta St. J. Bombay Univ. 'Kill, 

Nicolas, G. (1932). Association des Bryophytes avec d’autres Organismes. In Manual 
of Bryology, edited by Fr. Verdoorn, Hague, pp. 109-28. 

Du Plessis, S. J. (1933). The life history and morphology of Olpidiopsis Ricciae nov.sp. 
infecting Riccia sp. in S. Africa. Ann. Bot., Bond., xlvii, 755-62. 

Shanor, Leland (1940). Studies in the genus Olpidiopsis. III. Some obser\^ations on 
the host range of certain species. J. Elisha Mitchell Sci. Soc. lvi, 165-76. 

Sparrow, F. K. (1931). Two new Ghytridiaceous fungi from Gold Spring Harbor. 
Amer. J. Bot, xviii, 615-22. 

Srinivasan, X. S. (1939). An ascomycetous fungus attacking Himalayensis St, 

Trans. Brit Myc. Soc. xxin, 55-62. 

Whiffen, Alma j. A. (1942). A discussion of some species of Olpidiopsis and Fseudo -- 
olpidium. Amer. jf. Bot xkix, 6 oy-ii. 

Whiffen, Alma J. A. (1943). New species of Nowakoskiella and Blastocladia. J. Elisha 
Mitchell Sci. Soc. lix, 37-43. 

[Accepted for publication 10 October 1946) 

[ 13 ] 


By C, H. GADD and C. A. LOOS, Tea Research 
Institute of Ceylon 

(With Plate I and 26 Text-figures) 

Many beetles of the family Scolytidae have long been known to live in 
association with specific fungi, the so-called ^ambrosia’, which form the 
main, and possibly the sole food supply of their larvae. Yet little is known 
of these fungi, as may be seen from the review of the literature given by 
Leach, Hodson, Chilton & Christensen (1940). 

XyleboTus fornicatus Eich., the shot-hole borer of tea, is an important 
ambrosia beetle pest in many tea districts of Ceylon, because the female 
beetles make galleries in living tea stems, which in consequence tend to 
break rather easily and so cause loss of crop. The beetle also invades other 
plants, of which the most important is the castor-oil tree. The beetle from 
tea is smaller than that from castor, measuring about 2^3 mm. in length 
as compared with 2*5 mm. from castor, and its elytral curve is more 
convex from scutellum to sutural apex. Because of these morphological 
differences, which, however, are not constant, Eggers (1922) described 
specimens from tea as a new species, X, fornicatior. Sampson (1923) and 
Beeson (1925) were unable to X, fornicatior zs a valid species, 

though Beeson considered the retention of the name in subspecific rank to 
be desirable on biological evidence. Beeson (1941) stated that in north- 
east and north-west India and in Indo-China the tea bush is rarely 
attacked hy X, fornicatus, although the borer is abundant in other hosts in 
the same locality, and there is no trace of the fornicatior characters. In 
Ceylon (Gadd, 1946) experiments have demonstrated that the beetle 
from castor will bore into tea stems and the beetle from tea into castor 
stems, but whether the host plants are normally interchangeable in nature 
seems doubtful. Nor, by any means, do the beetles from tea always show 
fornicatior characters ; they are more often indistinguishable from castor 
beetles. For this reason the specific name will be used here 

throughout to avoid any implication that galleries in tea stems had been 
made by beetles with yomVflifw characters. 

The shape and size of galleries vary to some extent with the size of the 
branch in which they are made. Tea stems about half an inch in diameter 
and less seem to be preferred, whereas castor stems are usually much 
thicker when attacked. Galleries are made by females alone without any 
assistance from males. Each starts with a straight entrance gallery of 
variable length running radially into the stem. In small tea stems the 
entrance gallery stops at the pith, along which a straight longitudinal 
gallery is then made. More frequently, especially in thicker stems, the 
entrance gallery terminates in the wood near the camt)ium and the branch 

14 Transactions British Mycological Society 

galleries are circular, running parallel with the outer surface of the stem, 
often very close to the cambium yet not damaging it. All galleries are 
communal, there being no special brood cells or chambers. Eg:gs are 
usually found near the gallery ends, often in small heaps of five or six, but 
larvae, pupae and adults may be located in any part of a gallery. 

Ambrosia can be seen as a fine, white, frost-like dust on the gallery wall, 
particularly where young larvae are feeding. It is not usually visible on 
the dark-stained walls of old galleries; nor does it ever occur in such 
quantity as to block the passage. There is no carefully prepared bed or 
layer of chips and excreta for the fungus to develop on, as mentioned by 
Imms (1934, p. 531) in his short general account of ambrosia, nor can 
two superimposed fungal layers be distinguished as described by Trotter 
(1934) from galleries made by an unnamed beetle in branches of living 
Brownea grandiceps. 

Sections show that the fungus consists of a thin surface layer from which 
arise numerous short, erect, unbranched, septate conidiophores, at the 
ends of which are solitary club-shaped spores, aseptate at first but with 
several (up to five) transverse septa later (PI. I, fig. i). This fungus 
belongs to the genus Monacrosporium. Amongst the conidiophores, close to 
the gallery walls, a few globose cells about 30 ja in diameter have been 
observed in tea stems, but not in castor. When stained, these globose cells 
each contain what appears to be a two-celled body (Text-figs, i, 2). The 
nature of these cells is not understood and they do not occur in pure 
cultures of xh.^ Monacrosporium. In galleries in castor the conidiophores are 
somewhat more septate and the spores shorter and broader, but in other 
respects the fungi appear ahke. Spores from galleries in tea (Text-figs. 3-5) 
range from 34-51 x 10-5-13 p, while those from castor (Text-figs, ri, 12) 
range from 25-37 x Mean measurements of fifty spores were 

40 X 1 1-5 ff om tea, and 32 x 13 /x from castor. 

Speyer (1918), in a short statement on the fungi from shot-hole borer 
galleries in tea stems, wrote : ‘ Observations on the fungus upon which the 
larvae of shot-hole borers feed show that two fungi are generally present 
in healthy galleries. These have been identified by Mr Fetch as Mona- 
crosporium and a conidial ambrosia fungus. The spores of the former appear 
in the gallery about seven days after the beetle has entered the branch. 
It is known that the conidial stage of some ambrosia fungi are modified 
Stages of fungi which grow free, on certain media, in a totally different 
form; there is therefore a possibility, though a remote one, that Monacro- 
sporium the free-living phase of the ambrosia grown by shot-hole borer.’ 
In a later publication (1923) he gave another description: ‘The ambrosia 
mycelium grown by X. fomicatus consists of comparatively long, sparsely 
septate, narrow hyphae; curiously enough, the rounded conidial bodies 
characteristic of other ambrosia fungi have not been observ-ed. In 
addition, spores of Monacrosporium are constantly found in the tunnels 
when this insect is excavating the gallery, and this fungus occurs also 
amongst the ambrosia hyphae.’ 

Speyer was evidently of the opinion that two fungi occur in the galleries, 

though how they can be distinguished in the absence of ‘rounded" conidial 
V - 

The ambrosia Fungus of ^ylthoxus fomic2L^ 15 

bodies characteristic of other ambrosia fungi’ is not apparent. Whether 
the globose cells amongst the conidiophores previously referred to in 
galleries in tea belong to a distinct fungus has not been determined, and 
they have not been observed in cultures of Monacrosporium. The available 
evidence indicates, however, that they are of no vital importance in insect 

If by the term ambrosia is meant the actual food of beetle larvae, then 
the Monacrosporium spores are ambrosia. The spores can be found in con- 
siderable numbers in the anterior gut of larvae freshly removed from 
galleries, and newly hatched larvae have been brought through all stages 
to pupation on pure Monacrosporium cultures derived from castor galleries 
in tea and castor. Nothing other than the fungus appears essential for 
their growth. . 

Single-spore cultures were easily obtained by plating spores from freshly 
opened galleries in tea or castor stems. The spores germinate rapidly, 
aseptate spores becoming septate and forming germ tubes within four 
hours. Germ tubes are normally produced at the apical and basal cells 
though they may arise from any cell (Text-figs. 6~8, 13-16). The fungus 
grew well on both malt agar and Waksman’s agar without acid ( Waksman, 
1927, p. 19) at room temperature at the St Coombs Laboratory (64-76^ F.). 
These were the only culture media tested, but it is probable that others 
would be equally suitable. 

The cultures obtained from spores in galleries in tea were easily dis- 
tinguished from those obtained from castor by their slower growth and 
greater colour development. At 26° C. cultures from tea, on plates, grew 
at an average rate of rather more than 5 mm. per day, whereas the castor 
cultures grew almost 9 mm. per day. Castor cultures were distinctly zoned 
(PI. I, fig. 3) and had regular smooth edges, whereas tea cultures were 
less distinctly zoned with broken, rugged edges, and were markedly 
coloured (PL I, fig. 2). In tubes, the aerial mycelium of castor cultures 
was more cotton-woolly than those from tea, which had a looser and 
shorter* mycelium. The colour change in plate cultures from tea became 
noticeable about the fourth day when the aerial mycelium near the centre 
of the colony became flesh coloured. Later the colour deepened to 
salmon, but the growing edge remained white. This colour development 
was not noticeable in tube cultures, nor in any cultures from castor. The 
agar of tea cultures, particularly slants, becomes deeply stained, dark 
vinaceous, near Corinthian purple (Ridgway, pi. 38), whereas in castor 
cultures the colour never attained a similar depth; it approximated 
Daphne red (Ridgway, pL 38). 

It may be appropriate to mention here that the wood adjacent to 
galleries in tea stems is usually stained dark purple (Speyer, 1922), though 
some variation occurs both in the colour and amount of it. The wood 
around galleries in castor is, however, rarely coloured. These observations 
support VerralFs (1943) statement that the stain around beetle tunnels is 
caused by the ambrosia fungi, which may easily be isolated from such 
wood. He attributed the intensification of colour to reactions of these 
fungi with insect secretions and other organisms, but in the writers’ 

l6 Transactions British Mycological Society 

opinion it may be due largely to the strain of the ambrosia carried by 
the beetles. 

In microscopic detail very little difference was to be observed between 
the two cultures. The spores are normally produced on short unbranched 











Text-fig. ; 

Text-figs. 1-26. Monacrosporium ambmsium n.sp. 

5. r, 2. Globose bodies from galleries in tea stem. 

5. 3-5, Mature spores from tea. 

5.6-8. Germinating spores from tea. 

9 > Immature spores and conidiophores from culture (tea) 

!. II, 12. Mature spores from castor. 

!. 13-16. Germinating spores from castor. 

17-19. Mature spores from culture (tea). 

. 20,21. Mature spores from culture (castor). 

22. Germinating spore from castor culture. 


.24,25. Spores and conidiopbores from castor gallery 
20, Immature spores in tea gallery. 

conidiophores, though in tubes in which larvae had fed branched conidio- 

with a 

dium (1 ext-hg. 23) . The conidiophores are very variable in length usually 
much longer and less septate than those observed in gallerms ’ Spores 
varied somewhat m shape, but were generally clavate^with 2-5 septa 

The ambrosia Fungus of Xyleboms fomicatus 17 

The spores from tea were not noticeably larger than those from castor, 
though fifty measurements of each gave the following results: tea 28-43 x 
8-12 (mean 36 x 9*5 jjl) ; castor 22-40 x 7-10*5 (mean 33 x 8*5 

Spore production was markedly stimulated by removing aerial mycelium 
with a sterile bent needle. After twenty-four hours a large number of 
small pustules of spores appeared on the agar surface. Aerial mycelium 
did not again grow from the cleared surface, but the pustules or globular 
masses of spores increased in size (PL I, fig. 4). Cultures from tea and 
from castor behaved alike in this respect. The same thing probably occurs 
in the galleries. The larvae feed mainly, if not solely, on spores, as spores 
alone have been found in the gut of larvae examined. Their feeding 
evidently stimulates spore production, for it is in those parts of a gallery 
where larvae are feeding that spores can be found in large numbers. 

In feeding experiments larvae hatched from eggs on damp blotting 
paper have been transferred to tea and castor cultures. Larvae from both 
sources fed readily on both cultures, and it appeared immaterial whether 
the tea or castor ambrosia was used as food supply for either beetle. 
Larvae were successfully brought to the pupal stage on both cultures. 

Experiments clearly demonstrated that the food of Xyleboms fomicatus is 
the fungus Monacrosporium described. How the parent beetle carries it 
from the original gallery and plants it in her new gallery is not known. 
Several hypotheses have been advanced, but they need not be considered 
here. The importance of the fungus in the insect’‘s economy, and the 
amount and nature of the work to be done before it can be planted in a 
suitable place for growth, seem to rule out the possibility that transfer is 
entirely accidental. But whether the transfer is accidental or due to 
instinctive actions, there can be little opportunity for the intermixing of 
the fungi carried by different families. It would not be surprising therefore, 
if numerous strains of the fungus exist, differing from one another in minor 
characters, such as colour development, determinable in comparable 
artificial culture. For this reason we regard the differences observed in 
culture between the two strains used in these studies as of minor im- 

So far as we are aware, this fungus has not been described previously; nor 
do we know of another ambrosia fungus belonging to the genus Monacro- 
sporium. We therefore propose for it the name Monacrosporium ambrosium. 

Monacrosporium ambrosium n.sp. 

The ambrosia fungus of the beetle Xyleborus fornicaMs Eich. Vegetative 
hyphae in the wood adjacent to the galleries and thinly lining the gallery 
walls, thin, hyaline, septate and branched. Conidiophores produced on 
the gallery walls, crowded, erect, septate, normally unbranched, rarely 
exceeding the spore in length. Spores solitary, terminal, hyaline, clavate, 
non-septate at first, with rounded apex, later and before germination 
becoming 2“5 septate and with a bluntly pointed apex, 25-51 x i 0*5-1 5-5 jx. 

Habitat. In galleries oi Xyleboms fomicatus Eich. in Camellia sinensis [ttdi) ^ 
Richms communis (castor-oil tree) and other stems invaded by the 


1 8 Transactions British My cological Society 

Hyphae steriles in ligno contiguo, etiam in parietibus internis cunicu- 
lorum effusae^j tenues, hyalinae, septatae, ramosae. Gonidiopliora iii 
cuniculis eiFormata, dense congregata, erecta, septata, plerumque Iiaiid 
ramosa, longitudine rare sporam excedentia, apice monospora. Goiiidia 
hyalina, clavata, prime continua apice rotundata, demum ante germiiia- 
tionem 2-5 septata, in apicem obtusum angustata, 25-51 x iO'5 — 15*5/^. 

Hab. in cuniculis Xylebori fornicati Eich., in caulibus CameMiae sinensis , 
Ricini communis et arborum aliarum. 

We are indebted to Miss E. M. Wakefield, M.A., F.L.S., of the 
Herbarium, Royal Botanic Gardens, Kew, for the Latin diagnosis. 


The fungus commonly associated with the ambrosia beetle Xjkborus 
fornicatus Eich., the shot-hole borer of tea and other plants, is described 
as new, and named Monacrosporium ambrosium. This fungus is undoubtedly 
the main, jf not the sole, food of the beetle, as larvae have been brought 
from the egg to pupation on cultures of the fungus. 


Beeson, C. F. C. (1925). Xylebortis fornicatus in India. Trap. Agrimiiuristy Lxv, 371-2. 
Beeson, C. F. C. (1941)- The Ecology and Control of the Forest Insects of India md the JVeigh- 
bouring Countries. Dehra Dun. 

Eggers, I. H. (1922). Kulturschadliche Borkenkafer des indischen Archipels. EnL Bet. 

Amst. VI, no. 126, pp. 84-8. Abstract in Rev. appL Entom. A, x, 572. 

Gadd, G. H. ( 1 946) . Studies of shot-hole borer of tea. i . Distr ibution and iionienclat lire. 
Tea Qmrt. ismii, 46-54. 

, Imms, a. D. (1934). A General Text Book of Entomology, 3rd ed. London. 

Leach, J. G., Hodson, A. C., Chilton, St J. P. & Christensen, M. (1940). Obser- 
vations on two ambrosia beetles and their associated fungi. Pkytopatk. xxx, 227-3(>» 
Sampson, F. W. (1923). Notes on the nomenclature of the family Scolytidac. Ann. Mag. 

nat. Hist, xi, no. 62, pp. 269-71. Abstract in Rev. appL Entom. A, xi, 257. 

Speyer, E. R. (1918). Progress repoi't on investigations into shot-hole borer of tea* 
Trop. Agriculturist, l, 373-4. 

Speyer, E. R. (1922). Shot-hole borer of tea : Damage caused to the tea bush. Bull. 
Dep. Agric. CeylonyUO, 60. 

Speyer, E. R. (1923). Notes upon the habits of Ceylonese x'-\nibrosia beetles. Bull. enL 
Res. XIV, 11-23. 

Trotter, A. (1934). II fungo-ambrosia delle gaiierie di un Xyleborino di Oyion. 

Ann. 1 st. sup. agr. Portici, s Ser. vi, ,256-75. Abstract in Rev. App. Myc. xiv, 167. 
Verrall, a. F.' (1943). Fungi associated with certain ambrosia beetles. J. agrk. Res. 

LXVI, 135. 

Waksuan, S, A. Principles of Soil Microbiology. London. 


Fig. I . Section through gallery in tea stem (pith) showing spores, x 280. 

Fig. 2. Culture on Waksman’s agar 8 days old, tea strain, x | . 

Fig. 3. Culture on Waksman’s agar 8 days old, castor strain, ’"x .L 
' ■ . ■ '■ '■ 4 *; ■' Spore pustules ' produced, after .removing aerial myceiium from culture, x 30* 

{Accepted for publication 15 October 1946) 

[ 19 ] 


By T. fetch 
(W ith Plates II-IV) 

In their general account of the fungi of Ceylon (J. Linn. Soc. xiv, 31), 
Berkeley and Broome wrote ^ Mar asmii and Lentini are, as might be ex- 
pected, abundant , and in their systematic list they enumerated forty-four 
species of Marasmius.^ thirty-five of them new species. Unfortunately, only 
thirteen of these were illustrated in the series of paintings of agarics, sent 
with s|>ccimens by Ihwaites and now at Peradeniya, from which Berkeley 
and Broome described most of their Ceylon species. There are also at 
Peradeniya specimens of Ceylon fungi which were erroneously supposed to 
have been named by Berkeley and Broome, cotypes in fact, but a later 
examination of some T-hwaites correspondence showed that they had not 
been seen by them. As with other cryptogams, Thwaites divided each of 
his gatherings into two parts, one of which was sent to Berkeley and Broome, 
while the other was retained at Peradeniya, mounted on paper duly 
numbered, the names being added later after the publication of the 
Fungi of Ceylon b 

Thwaites collected his specimens over a fairly long period, and he 
placed under the same number those he thought were the same species. 
Naturally in some instances be was mistaken, and Berkeley and Broome 
divided up his specimens if they thought that was so. For example, 
Thwaites’s no. loi was collected on four dates ranging from November 1867 
to Septem!>er 1868, and Berkeley and Broome divided it into four MarasmU 
and a Colljfm. But there was only one figure, and one can only surmise 
that the other species resembled it sufficiently closely to make Thwaites 
think that they were the same. It proved impossible to sort out these and 
similar mixtures from the specimens at Peradeniya, and the revision of the 
Ceylon MarasmU had to be postponed until Berkeley’s specimens in Herb. 
Kew. and Broome's specimens in Herb. Mus. Brit, (not always the same) 
could be examined. Specimens were collected, described, and tentatively 
named, and during my home leave in 1911 and again in 1915, some pro- 
gress was made in their determination, but with the expansion of the work 
of' the mycokigical division this project ultimately lapsed. 

The following include redescriptions of most of Berkeley and Broome’s 
Ceylon species. The regions covered were practically those in which 
Thwaites collected his mycological specimens, but even these have not 
been exhausted. The low country, in both the wet and the dry zones, has 
yet to be investigated. 

Alarasmius nephelodes (B. & Br.) Petch, Ann. Perad. iv, 403; Agaricus 
{Collyhia) nepheiades B. & Br., Fungi of Ceylon, m. 105; Marasmus ockraceu^ 
B. & Br., Fungi of Ceylon, no.' 355. ; ■ ) 

Pileus broadly convex, centre even or irregularly depressed, reddish 
brown in the centre, ochraceous elsewhere, sometimes feebly zoned, 

52 -'S 

20 Transactions British Mycological Society 

margin sometimes whitish, fleshy, smooth, slightly hoary when youn^ 
shining, 2-4 cm. diameter, flesh white, becoming reddish when cut; stalk 
4-5-5 cm. long, 3-4 mm. diameter, nearly equal, white, sometimes longi- 
tudinally streaked ochraceous, becoming reddish or ochraceous when 
handled, pruinose or minutely tomentose, tough, hollow, base tomentose; 
gills pallid, distant, free or slightly adnexed, abruptly truncate behind; 
spores white, narrow oval, On the ground among grass, 
solitary or in groups. Frequent on the lawns at Peradeniya (PI. II, fig. i). 

Gardner’s painting no. 70, referred by Berkeley to Hjgrophorus obrusseus 
Fr., resembles this species, but as it was said to grow on the ground in 
woods it may be something else. 

Marasmius calvus B. & Br., Fungi of Ceylon, no. 354. 

Pileus up to 4-5 cm. diameter, hemispherical then broadly convc.x, 
sometimes slightly umbonate, fawn-coloured to ashy, bay brown in the 
centre, minutely pruinose, irregularly radially striate, becoming sodden 
and dark brown at the margin and along the striae, flesh white, thin ; stalk 
up to 8 cm. long, 4 mm. diameter, white, longitudinally striate, often 
twisted, glabrous, white tomentose at the base, brittle, stuffed then hollow, 
brown internally; gills pallid, broad, distant, rounded behind, adnexed. 
On the ground among dead leaves. No. 41 15, Hakgala, 28 September 1914 
(PL I, fig. 8). 

Berkeley and Broome described this species as ‘umber, elegantly varied 
with radiating lines’, and the gills as ‘sometimes ventricose, as in the more 
beautifully painted specimens’. Thwaites’s no. 766, however, was a mixture, 
both as regards specimens and paintings, and the figure elegantly varied 
with radiating lines is of a young specimen of M. rivulosus. 

Marasmius purpureo-griseus Petch, n.sp. 

Fasciculate. Pileus broadly convex, purple grey, paler towards the 
margin, not sulcate, minutely radially silky, about 1-5 cm. diameter, 
margin incurved, flesh white, thin; stalk up to 4 cm. long, 1-5 mm. 
diameter, brownish white, shining, cartilaginous, stuffed then hollotv, 
fistulose; gills crowded, white or cream-coloured, broad, adnatc or 
sinuato-adnate, edge straight. On dead wood, attached by white myce- 
lium. No. 5736, Henaratgoda, 25 June 1918. The specimen became 
rather dry, before the painting could be made (PI. Ill, fig. 7}. 

Marasmius muhijugus (B. & Br.) Petch, in Trans. Brit, mycol. Soc. xxvii, 138; 
A. {Collybia) multijugus B. & Br., Fungi of Ceylon, no. too. 

Usually densely caespitose, sometimes connate at the base or up to half 
the height; pileus conico-campanulate, apex rounded, margin recurved 
when old, or convex or almost plane, centre sometimes depressed or almost 
umbilicate, up to 5 cm. diameter, ashy to red brown, glabrous when moist, 
innately silky when dry, irregularly plicate half-way to the centre, flesh 
very thin; stalk up to to cm. long, 7 mm. diameter, connate at the base, 
sometimes forking, often flattened and twisted, white, smooth or minutely 

Revisim of C^lon }Aax2LSvim. T. Fetch 21 

pruinose, equal, hollow, cartilaginous; gills pallid, becoming brown or 
liver-coloured from the edge upwards, distant, equal, rather broad, 
rounded behind, adnate or almost free, interstices veined; spores white, 
narrow oval, inequilateral, or somewhat attenuated to the apiculus, 
apiculus prominent curved, S-gxg-qja. Smell resembles that of M. 
oreades. On the ground, often with coarse rooting mycelium. Peradeniya, 
6 April 1907; 30 June 1908; no. 6526, i May 1921 (PI. Ill, fig. 15). 

Marasmius leucophaeus (B. & Br.) Petch, in Trans. Brit, mycol. Soc. xxvn, 138; 
A. {Collybia) leucophaeus & Br., Fungi of Ceylon, no. 113; Marasmius 
cornicolor B. & Br., Fungi of Ceylon, no. 361. 

Generally densely caespitose, sometimes scattered. Pileus usually 
2-3-5 cm. diameter, plane, margin often repand, feebly striate in wet 
weather, centre even or depressed or umbilicate, or broadly convex, up to 
5-5 cm. diameter, irregularly plicato-striate almost to the centre, deep 
purple brown, sometimes becoming paler outwards, often with an 
abruptly paler margin on small specimens and sometimes with a pale 
central spot over the stalk, appearing glabrous but covered with minute 
adpressed hairs, drying to ashy brown and minutely hoary, cartilaginous, 
thin, flesh purple brown; stalk up to 5 cm. long, 1-2 mm. diameter, dark 
purple brown when moist, ashy brown when dry, equal or slightly 
attenuated upwards, glabrous or minutely tomentose, often with a white 
tomentose ring at the apex, stuffed then hollow, often connate, base some- 
times tomentose; gills narrow and crowded on small specimens, rather 
broad and subdistant on larger, adnate to the expanded apex of the stalk, 
pallid, becoming brown from above downwards, often retaining a pale 
edge; spores white, narrow oval, 6-10 x 3-4/a. On the ground among 
dead leaves, etc. Common at Peradeniya (PI. Ill, fig. 2). 

A form occurs on the rough bark of living trees, e.g. jak, which appears 
very different, except during heavy rains, when it assumes the normal 
form. It is usually scattered, pileus at first hemispherical, then plane, up to 
I cm. diameter, grey, purple grey, or bluish grey, hoary with adpressed 
fibrils and arid-looking. This form is no. 3265, 13 December 1911; no. 
4172, II October 1914; no. 4206, 25 October 1914, all on jak trees, 

The specimens in Thwaites’s no. 204, noted by Berkeley and Broome as 
dull reddish purple to fuscous, and assigned to M. Wynnei var. auroricolor in 
Fungi of Ceylon, no. 353, are M. leucophaeus. 

Marasmius rivulosus B. & Br., Fungi of Ceylon, no. 372. 

Pileus campanulate to broadly convex, broadly umbonate, often de- 
pressed round the umbo, radially corrugated almost to the umbo, pale 
brown or tan, the colour varying with age and weather conditions, some- 
times dark brown in the centre, ashy when dry, glabrous, about 4 cm. 
diameter but sometimes up to 10 cm., flesh white, thin except over the 
stalk; stalk up to 9 cm. long, 3 mm. diameter, base slightly expanded, 
longitudinally striate, at first smooth and shining, becoming finely white 

22 Transactions British Mycological Society 

tomentose, pallid then purple brown, fibrous in wet weather, subcarti- 
laginous when dry, stuffed then hollow, internally brown, often excentric j 
gills white or pinkish, becoming brown, moderately crowded, broad near 
the stalk, elsewhere narrow, arcuate, adnate or adnato-decurrent, con- 
nected by veins; spores white with a yellow tinge, varying from oval, 
6 x4/x, to clavate or narrow oval with a thick excentric apiculus, lo- 
13 x5^, in the same spore print. On decaying stumps or on a felted 
mycelium on chips of wood, scattered or clustered ; stalks usually excentric 
when growing on stumps. Common at Peradeniya (PI. II, fig. 2). Young 
specimens are sometimes purple brown at first, the colour changing to 
pale brown first in the centre, leaving the margin darker brown or brown 
in the furrows. 

Specimens on a living palm stem {Wallichia), no. 6626, 12 June 1923, 
and on coir in orchid pots, no. 6762 a, 19 June 1924, are more delicate and 
subpellucid, but otherwise the same. 

Marasxnius rufo-ochraceus Petch, n.sp. 

Fasciculate or scattered. Pileus broadly convex to almost plane and 
irregularly undulating, up to 3-5 cm. diameter, red brown in the centre, 
becoming ochraceous or pallid outwards, or wholly ochraceous when dry, 
irregularly radially grooved, glabrous, thin, cartilaginous; stalk up to 
3 cm. long, usually compressed, up to 3 mm. broad, cartilaginous, red 
brown, becoming paler upwards, glabrous or very minutely tomentose, 
stuffed then hollow; gills moderately distant, slightly ventricose, adnate, 
pallid becoming red^sh brown, interstices strongly veined when old ; spores 
white, clavate, 10-14 x 5-6 /x.. On dead wood, Hakgala, no. 5583, 31 
December 1917; no. 5603, December 1917; no. 6653, 7 September 1923. 

Marasmius congregatus Mont-., Guy. no. 307; Cantkarellus elegans B. & Br., 
Fungi of Ceylon, no. 346; Marasmius pellucidus B. & Br., Fungi of Ceylon, 
no. 359. 

In dense clusters among dead leaves or on rotten wood, with copious 
white rooting mycelium. Pileus frequently angularly campanulate at 
first, i.e. with a flat top and more or less perpendicular sides, then broadly 
campanulate, rarely plane, usually umbilicate, rather closely radialK' 
rivulose up to the centre, glabrous, translucent, papyraceous but tough, 
margin undulating, often repand, generally about 4 cm. diameter but 
sometimes up to g cm., white or yellowish; stalk up to 9 cm. long, 2-3 mm. 
diameter, rigid, cartilaginous, hollow, straight or twisted, glabrous, equal 
or expanding upwards, dark purple brown at the base, paler upward.s and 
white at the apex, becoming entirely purple brown; gills white, usually 
narrow, about i mm. broad, but sometimes up to 4 mm., arcuate, forking, 
adnate to the expanded apex of the stalk, edge rather acute; spores %vhitc, 
clavate, 6-8 x 3 p. Common at Peradeniya (PL III, fig. i). 

Berkeley and Broome’s specimens of Cantharellus elegans were small 
examples; after the description they added ’’ Marasmius congregatus Mont.'. 
Saccardo interpreted that as ‘hue spectat M. congregatus". Evidently 
Berkeley and Broome regarded the latter name as a synonym. 


Revision of Ceylon Marasmii. T. Fetch 

Marasmius crispatus B. & Br., Fungi of Ceylon, no. 357. 

Clustered or scattered, among dead leaves. Pileus up to 7 cm. diameter, 
broadly convex, then plane, or repand and undulating, centre umbilicate, 
irregularly radially plicate almost to the centre, thin, tough, at first pallid 
grey brown, becoming buff on drying, glabrous, appearing cartilaginous 
when moist, but innately fibrillose when dry; stalk up to 9 cm. long, 4 mm. 
diameter, often compressed and up to 7 mm. broad, equal, expanding at 
the apex, cartilaginous, stuffed then hollow, dull, minutely powdered at 
first, becoming minutely tomentose, clothed with white tomentum at the 
base, with a mealy or minutely tomentose zone at the apex, coloured as the 
pileus at first, becoming purple brown from the base upwards; gills distant, 
arcuate, narrow, up to 4 mm. broad, sometimes forked, attenuated out- 
wards, rounded behind and adnate to the expanded apex of the stalk, 
coloured as the pileus; spores white, oval, inequilateral, 8-10 x 4 /x. No. 
4239, Hakgala, 30 October 1914; no. 5235, Peradeniya, July 1917 (PL II, 
fig. 6). 

In Herb* Peradeniya, there are two species in the specimens and paintings 
marked no. 766, if. calvus. That dated November 1868 is if. crispatus, 
Thwaites also sent specimens of the latter with if. congregatus in no. 38. 
It looks like a coarse if. congregatus, 

Marasmius Caryotae (Berk.) Petch, A?in, Perad. iv, 403; Heliomyces Caryotae 
Berk., Hooked s Lond, J, Bot. (1847)^ p, 491. 

Pileus 1-2*5 diameter, rarely up to 5 cm., campanulate, occasionally 
depressed in the centre, then expanded with a recurved margin, deeply 
silicate almost to the centre, glabrous, somewhat coriaceous, thin, pale 
yellow or greyish yellow, becoming ochraceous or pale brown when old, 
red brown when dried; stalk 4-8 cm. long, 1-2 mm. diameter, white 
becoming yellow, brown when old, rigid, hollow, almost glabrous, 
cartilaginous, tomentose at the base; gills distant, broad, somewhat 
ventricose, adnexed, rather thick, greyish yellow, brown when old; spores 
white, narrow oval or clavate, rather thick-walled, 20-25 x 5 /x. On the 
ground among grass, usually in troops ; common on the lawns at Peradeniya 
(PL II, fig. 3). ■ 

Marasmius hypochroides B. & Br., Fungi of Ceylon, no. 356. 

Pileus up to 3 cm. diameter, broadly campanulate, margin irregular in 
expanded specimens, rather pale red brown or olive brown, very thin but 
tough, not sulcate, cells of the pellicle crowned with close-set conical spines 
up to lOjLx long; stalk cartilaginous, hollow, fistulose, terete, 2 mm. 
diameter, or flattened, up to 4 mm. broad, equal or attenuated upwards or 
expanded upwurds, glabrous, red brown below, yellow brown above; gills 
white, sometimes yellowish, distant, ventricose and up to 6 mm. broad in 
large specimens, free, interstices veined; spores white, obliquely oval, 
8-1 3 X 4-6 ju. Among dead leaves, etc. Peradeniya, no. 4208, 25 October 
1914; no. 4244, 4 November 1914; no. 4372, 13 December 1914 (PL IV, 
figs. 10, 1 1). 

Berkeley and Broome described the pileus as deeply sulcate; it has that 

24 Transactions British Mycological Society 

appearance in the dried specimens, because of the tenuity of the pileus and 
the thickness of the gills, but fresh specimens are even, becoming sulcato- 
striate when collapsing. 

Marasmius bruimeostriatus Fetch, n.sp. 

Pileus campanulate, then almost plane, obtusely umbonate, up to 2 cm. 
diameter, rufous, centre darker brown, margin paler, yellow brown, 
radially sulcate, deeper coloured in the sulcae, minutely rugose, margin 
often repand; stalk up to 2 cm. long, i mm. diameter, expanded at the 
apex, cartilaginous, red brown, pruinose or minutely tomentose, base 
tomentose, arising from a thin stratum of white mycelium; gills pale 
ochraceous, distant, rather thick, narrow, equal, adnate; spores white, 
clavate or oval, 5-9 x 3-4 fx. On dead leaves and twigs. No. 4116, Hakgala, 
29 September 1914 (ft. 11 , fig. 7). 

Marasmius fulviceps Berk., Hooker's Lond. J. Bot. (1847), p. 490; Marasirlms 
nummularius B. & Br., Fungi of Ceylon, no. 351. 

Pileus hemispherical, then broadly convex or almost plane, pale yellow 
or pale ochraceous, deeper ochraceous in the centre, or yellow brown, or 
sometimes deep brown, up to 1*5 cm. diameter, slightly wrinkled in the 
centre, radially striate elsewhere, glabrous; stalk up to 4 cm. long, 
0*75 mm. diameter, blackish brown at the base, then red brown, yellow 
brown or white at the apex, cartilaginous, hispid, strigose at the base; gills 
crowded, narrow, white then pale ochraceous, adnate, the gill tissue 
continued from gill to gill round the apex of the stem; spores white, 
narrow oval, inequilateral, 10-12 x 4 fc. On decaying leaves and twigs, 
scattered or clustered, arising from a sheet of tawny mycelium, frequently 
with innate strands radiating from the base of the stalk, Peradeniya, no. 
2903, 9 August 1909; no. 3505, 14 July 1912 (PL IV, fig. 6). 

This species was sent from Ceylon by Gardner, with a figure. The 
Thwaites specimen referred to it by Berkeley and Broome (Thw. 807*) is 
M, haematocephalus, 

Marasmius pallidorubens (B. & Br.) Fetch, in Trans. Brit. mycoL Soc. xxvn, 
138; A. {Mycena) pallido-rubens B. & Br., Fungi of Ceylon, no. 1 19. 

Pileus campanulate or broadly convex, obtusely umbonate or depressed 
in the centre, membranous, translucent in wet weather, up to 4 cm. 
diameter, pale lavender, purplish brown in the centre, or in dry w’cathcr 
wholly vinouSj strigose in the centre, elsewhere covered with scattered 
small scales with red brown points, plicato-sulcate almost to the centre, 
margin slightly serrate with brown points, flesh thin; stalk up to 3 cm. 
long, 2*5 mm. diameter, equal or slightly attenuated upwards, colour of 
the pileus, becoming red brown, covered with minute red brown points, 
hollow, subtranslucent; gills . distant, ventricose, sinuato-adnate or 
adnato-decurrent, lavender becoming purplish red, edge clothed with red 
brown points, interstices veined; spores white, subglobose, or somewhat 
pyriform with an apical apiculusf 8-9 x 7-8 /x. The pileus and gills turn 
rather dark red when bruised; stains paper red. On dead wood. Pera- 

Revision of Ceylon Marasmii. T. Fetch 25 

deniya, no. 4277, 7 November 1914; no. 4382, 19 December 1914 (PL II, 
fig- 5 )- 

After the description, Berkeley and Broome stated ‘ no. 937 is possibly the 
same species. Spores -0003 by -000255 echinulate’. The later statement is 
entered in Saccardo for Mycena pallido-mbens, but no. 937 in Herb. Peradeniya 
is a diflferent species. 

Marasmius proximus B. & Br., Fungi of Ceylon, no. 368; M. obscuratus 
B. & Br., Fungi of Ceylon, no. 370, not of Fetch in Ann. Perad. vi, 56. 

Pileus at first hemispherical, then expanded, plane, or depressed in the 
centre with a decurved margin, sometimes with a broad convex umbo and 
a depressed ring round it, feebly sulcate almost to the centre when fresh, 
pale ochraceous, or ashy and brownish in the centre, about i cm. diameter, 
rugose, minutely scurfy when dry; stalk about 1-5 cm. long, 0-5 mm. 
diameter, red brown at the base, yellow brown above, becoming black 
brown, cartilaginous, minutely pruinose or tomentose, sometimes with a 
white tuft of hyphae at the base, especially when growing on dead leaves ; 
gills adnate or adnato-decurrent, pallid then pale brown, somewhat 
distant, rather ventricose. On dead leaves and twigs, sometimes accom- 
panied by a white glabrous film of mycelium. Peradeniya, no. 3359, 
January 1912; no. 4152, ii October 1914; no. 4205, 24 October 1914; 
no. 6553, 30 October 1922. Henaratgoda, nos. 5905, 5906, 25 June 1918. 
The pileus and gills sometimes turn red brown when dried (PL II, fig. 9). 

Berkeley and Broome cited Thwaites no. 93 in part for this species. The 
remainder of no. 93 was assigned to Naucoria furfuracea P., together with 
Thwaites no. 1182. The latter has been shown to be Flammula dilepis B. & 
Br. {Ann. Perad. iv, 53), and the remainder of no. 93 is Marasmius proximus. 
Maucoria furfuracea should be deleted from the Ceylon list. The type of 
Marasmius proximus in Herb. Kew. is marked 93 and 398. No. 398 was 
described as Omphalia delicia B. & Br.; I have not examined the type. 

The specimens and figure of Thwaites no. 804, which Berkeley and 
Broome placed as a variety of Marasmius subcinereus, are M. proximus. M. 
mutabilis was also Thwaites no. 804 (not 204) but the note ‘pale dun colour 
when fresh ’ refers to M. proximus, not to it. M. obscuratus B. & Br. was based 
on small specimens of M. proximus: 

Marasmius kirtelius B. & Br., Fungi of Ceylon, no. 382. 

Pileus at first broadly convex, then almost plane or with a decurved 
margin, centre depressed, often with a broad convex umbo and a depressed 
ring round it, feebly radially sulcate towards the margin when fresh, ashy 
or ashy brown, usually with a pink or purple tinge when fresh, sometimes 
with a pale central spot, surface of innate matted fibrils, appearing smooth 
or minutely lacunose, about i cm. diameter; stalk up to 2-5 cm. long, 
0-75 mm. diameter, cartilaginous, hispid, red brown, becoming black 
brown from the base upwards ; gills rather crowded, narrow, adnate, white 
or pallid, becoming pale brown; spores white, narrow oval, somewhat 
clavate, 6-8 x 3 /x. On dead leaves and twigs, sometimes accompanied 
by a white glabrous film of mycelium. Fairly common at Peradeniya; 

26 Transactions British Mycological Society 

no. 4152, 1 1 October 1914; no. 4294, November 1914; no. 430O5 November 
1914; no. 5807, 27 October 1918; no. 520^3 28 October 1918 (PI. II, fig. 43 
and PL III, fig. 13). 

Notwithstanding the figures, it is difficult to separate this species 
always from M. proximus. As a rule, the latter has broader and more 
distant gills. 

Marasmius mutabilis B. & Br., Fungi of Ceylon, no. 367 ; M. eximius B. & Br., 
Fungi of Ceylon, no. 371. 

Pileus infundibuliform, outer half sometimes almost plane, up to 2 cm. 
diameter, pale brown to reddish brown, glabrous, radially streaked with 
innate darker fibrils especially in the funnel, even, becoming radially 
sulcate, tough, thin, total height 1-5-2 cm.; stalk up to 1-4 cm. long, 
1-5 mm. diameter, expanded upwards, at first purple brown at the base, 
white above, becoming dark purple brown, cartilaginous, minutely 
tomentose, usually with a rather compact whitish cushion at the base, 
faintly longitudinally striate; gills pallid, becoming brown %vhen old, 
decurrent, rather distant, sometimes forked, arcuate, interstices veined; 
spores white, narrow oval, 8-9 X4p,. On dead wood and sticks. Pera- 
deniya, no. 2298, 12 August 1906; no. 4289, ii November 1914. 

This was part of Thwaites no. 804, not 204 as stated in the Fungi of 
Ceylon. Berkeley and Broome said it was analogous to Cantharellus rubidus. 
They probably meant C. rabidus, Fungi of Ceylon, no. 349, and Saccardo 
printed it as rabidus in the note to M. mutabilis. 

The type of ikf. eximius is a large specimen of M. mutabilis. It is infundi- 
buliform, with broad decurrent gills, and innate radial dark fibrils on the 

Marasmius senescens Fetch, n.sp. 

Pileus 1-5 cm. diameter, broadly convex, centre depressed, sometimes 
umbonate, even or irregularly plicato-sulcate, centre wrinkled, sordid 
grey with a paler margin, thin, centre slightly tomentose, elsewhere with a 
few scattered fibrils and here and there glistening particles; stalk 1 cm. 
long, 1-5 mm. diameter in the middle, expanding upwards, white or ashy, 
becoming black brown from the base upwards, compressed, striate, 
minutely longitudinally silky, cartilaginous; gills distant, adnate or 
adnato-decurrent, united behind over the stalk and separating, broad, 
slightly ventricose, rather thick, forked, pallid then brownish, interstices 
veined; spores white, narrow oval, somewhat pyriform, 7-10 x 3-4 /i. On 
the ground. No. 4306, Peradeniya, 15 November 1914. 

Aforajffzzaj ratebereaj B. & Br., Fungi of Ceylon, no. 369. 

Pileus convex, then plane, slightly depressed, or umbilicate, sometimes 
infundibuliform in wet weather, sulcate almost to the centre, thin, dark 
green when young, then greenish grey or greenish white, almost black in 
the centre, or grey in the centre, white on the outer half, blotched green, 
becoming greenish brown or pale greyish brown when old, up to 1-5 cm. 
diameter; stalk up to 2-5 cm. high, 0-5 mm. diameter, rigid, dull, bluish 

Revision of Ceylon T. Fetch 27 

black or black, pale towards the apex, hoary with minute white particles, 
solid, slightly thickened upwards, base either not thickened, or expanded 
into a disc up to 1*5 mm. diameter, or with a small tuft of white hyphae; 
gills white when young, then bluish or bluish grey, rather broad, distant, 
thick, adnate or slightly decurrent, interstices strongly veined; spores 
white, stellate, consisting of four broad-based triangular processes, each 
about 4 fi long and 3-4 fi diameter at the base, the distance from point to 
point being 8-12 /x. On young specimens, the pileus and gills turn blue to 
greenish black when bruised. On fallen branches and dead leaves. 
Peradeniya, not uncommon. No. 6627, 12 June 1923 (PL III, fig. 14). 

A form with a pure white pileus and gills occurred on the trunks of living 
nutmeg trees at Peradeniya. Pileus up to 1-5 cm. broad, orbicular, convex 
or almost plane; stalk excentric, curved, up to 3 mm. long, 0*5 mm. 
diameter, expanding slightly upwards, greenish black, or black below, 
bluish above, pruinose; gills white, arcuate in convex pilei, ventricose in 
plane examples; colour change when bruised, and spores, as in the type. 
No. 6542, 22 September 1922; no. 6709, 25 December 1923 (PI. HI, fig. 5). 

■Marasmius corticigena B. & Br., Fungi of Ceylon, no. 388. 

Pileus at first broadly convex, pale reddish brown in the centre, ashy or 
white towards the margin, margin even or faintly striate, minutely rugose ; 
then expanded, up to 2-5 cm. diameter, broadly convex or plane, pale 
reddish brown, irregularly sulcato-striate over two-thirds the pileus, centre 
innately radially silky; stalk at first white, becoming brown from the base 
upwards, finally black brown, purple brown at the apex, expanding 
above, minutely tomentose, cartilaginous, base slightly tomentose, up to 

2 cm. long, 0-8 mm. diameter in the middle; gills at first white, finally pale 
brown, subdistant, slightly ventricose, attenuated outwards, narrow, of four 
lengths, the longer adnate and united behind over the stalk, edges here and 
there tomentose with irregular, somewhat nodular hairs, up to 50 x8 ft, 
sometimes encrusted at the apex, interstices veined ; spores white, narrow 
oval or clavate, 8-12 x 3-5-"4ft. On living tree trunks. Peradeniya, no. 
4370, 13 December 1914; no. 6486, 19 July 1922 (PL 11 , fig. 10). 

Berkeley and Broome's specimens were quite immature, the pileus 

3 mm. diameter, margin incurved, and the stalk 5 mm. long, brown to 
black. They show the tomentum on the gill edges. 

.Marasmius niuosus' Berk,, Hookefs Land, Jf, Bot, (1856), p. 139- 

Pileus at first broadly convex, then plane, even, or umbonate in the 
centre, becoming depressed or subumbilicate, or with a ring furrow 
surrounding the umbo, irregularly radially sulcate almost to the centre, 
white, pale ochraceous in the centre, or pale ochraceous with a pinkish 
tinge, becoming ashy on drying, coriaceous, 1-2 cm. diameter, margin 
irregularly crenate; stalk concolorous with the pileus, but purplish iri very 
wet weather, up to 2 cm. long, i mm. diameter, equal, expanded slightly 
at the apex, cartilaginous, stuffed, glabrous or minutely pruinose; gills 
white, moderately crowded, arcuate, adnate or slightly decurrent. Among 

28 ■ Transactions British Mycological Society 

grass. Peradeniya, no. 6316, 20 November 1921; no. 6552, 30 October 
1922; no. 6624, 12 June 1923 (PL III, fig. 12). 

The fungus described above agrees with the specimens, Thwaites no. 
932, in the Peradeniya herbarium, identified by Berkeley and Broome, 
but not altogether with the original description. 

Marasmius stjpims B. & Br., Fungi of Ceylon, no. 389. 

Pileus hemispherical, up to 8 mm. diameter, sometimes expanded, 
broadly convex, up to 1-5 cm. diameter, even or obtusely umbonate or 
sometimes depressed in the centre, expanded specimens feebly radially 
sulcate, white, slightly ochraceous in the centre, pruinose, or minutely 
tomentose on unexpanded specimens, becoming minutely sprinkled witlx 
brown points or almost glabrous, brown when old ; stalk at first white, 
subtranslucent, rather stout, up to 1-5 cm. long, o-6 mm. diameter, 
becoming red brown from the base upwards and diminishing to 0-4 mm. 
diameter, finally entirely blackish brown, cartila^nous, dull, minutely 
fibrillose or pruinose; gills distant, white, becoming brown when old, 
adnate, arcuate, or somewhat ventricose on expanded specimens, edge 
serrate; spores white, 10-12 x4/x, clavate, apiculus obtuse and sublateral, 
making the spore sometimes appear hooked at one end. On sticks and 
bamboos : remarkable for the persistence of its juvenile condition. Pera- 
deniya, no. 2867, 4 July 1909; no. 2873, 18 July 1909; no. 2899, 5 August 
1909; no. 4248, 4 November 1914 (PI. IV, fig. 17). 

Marasmius stypinoides Petch, n.sp. 

Pileus white, greyish in the centre, broadly convex or almost plane, up to 
5 mm. diameter, centre umbilicate, feebly radially sulcate, minutely 
rugose, margin crenate; stalk up to 8 mm. long, usually curved, 0’4 mm. 
diameter, at first white, turning black from the base upwards, shining, 
minutely powdered, longitudinally striate, expanded at the apex; gills 
white, crowded, rather ventricose, the longer adnate to the expanded apex 
of the stalk and separating as if collared, edge serrate. On dead bark on a 
fallen tree trunk. Hakgala, no. 4029, April 1914. 

No. 4114, on a dead leaf, Hakgala, September 1914, is a similar species, 
but is umbonate. Unfortunately I have seen only one example. 

Marasmius cineraceus Petch, n.sp. 

Pileus infundibuliform, margin decurved, feebly radially sulcate, 9 mm. 
diameter, ashy, smooth, with a surface layer of interwoven hyphac; stalk 
black, with a white pruina, cartilaginous, i cm. long, 0-75 mm. diameter, 
equal, arising from a small circular glabrous disc ; gills moderately crowded, 
white, sigmoid, broad, adnato-decurrent, margin crenate, interstices 
veined. On a dead leaf. Peradeniya, no. 5589, October 1918. 

Marasmius Leveillianus (Berk.) Pat., Bull. Soc. Mycol. Fr. xxxiii (1917), 55; 
Heliomyces Leveillianus Berk., Decades of Fungi, no. 158; Marasmius um- 
braculum B. & Br., Fungi of Geylon, no, 365. 

Pileus up to 3-5 cm. diameter, at first hemispherical, becoming plane, 
umbonate, depressed round the umbo, radially sulcate, rigid when dry. 

Remsim of Ceylon Maxdsnm. T. Fetch 29 

deep red blown, glabrous; stalk up to 8 cm. long, 0-5 mm. diameter, black 
brown, equal, expanding suddenly at the apex, rigid, horny, glabrous, 
hollow; gills distant, usually all the same length, free, truncate behind, 
white becoming yellow, margin sometimes red brown, interstices sometimes 
veined; spores white or faintly yellowish, narrow oval, 8-10x3-4/1. 
Gregarious, on dead stumps, bamboos, etc., rather common. Peradeniya, 
18 June 1906; 15 July 1907; 15 July 1909, etc. Wariapolla, 6 December 
1906 (PI. IV, fig. 2). 

Marasmius atrorubens Berk., Hooker’s Lond. J. Bot. i, 188. 

Pileus campanulate at first, hispid with setae like those of the stalk but 
shorter, then broadly convex and glabrous or nearly so, i cm. diameter, 
deep red brown, somewhat mottled, margin faintly striate when fresh, 
grooved when dry, cells of the pellicle covered with short spines up to 5 p 
long ; stalk up to 4-5 cm. long, 0-5 mm. diameter, horny, rigid, red brown, 
brown when dry, hispid with brown, rigid acute setae, up to 120/1 long, 
3-4/1 diameter, apparently persistent; gills crowded, pallid then pale 
ochraceous, free. In groups among dead leaves, arising from yellowish 
strigose patches of mycelium. Peradeniya, no. 5176, 10 June 1917. 

The above description refers to Ceylon specimens which appear to fit 
Berkeley’s description. They have not yet been compared with the type 
from Cuba. The Ceylon specimens attributed by Berkeley and Broome to 
G- this species. 

Marasmius florideus B. & Br., Fungi of Ceylon, no. 380. 

Pileus campanulate or conico-campanulate, then expanded and margin 
repand, apex sometimes flattened, up to 2 cm. diameter, red brown, dark 
in the centre, becoming paler outwards, plicato-striate almost to the 
centre, thin but tough; stalk up to 5-5 cm. high, i mm. diameter, glabrous, 
horny, red brown to yellow brown below, pale at the apex, becoming 
blackish brown from the base upwards, base strigose; gills cream-coloured 
to yellowish, edge red brown, ventricose, adnate, moderately distant, of 
four lengths. Among dead leaves. Peradeniya, no. 4207, 25 October 1914 

(PI. IV, fig. 3 ). 

Marasmius confertus B. & Br., Fungi of Ceylon, no. 352 ; M. chondripes B. & 
Br., Fungi of Ceylon, no. 364; M. hemibaphus B. & Br., Fungi of Ceylon, 
no. 379. 

Pileus hemispherical, then expanded, broadly convex or almost plane, 
reddish brown in the centre, elsewhere tawny or golden brown, centre even 
or slightly depressed, margin feebly sulcate, usually about i -5 cm. diameter, 
exceptionally 3 cm., membranous, the cells of the pelhcle bearing close- 
set conical spines up to 6 /i long; stalk up to 3-5 cm. long, 0-5 mm. diameter, 
rigid, horny, shining, often flexuose, yellow brown, red brown at the base, 
white at the apex at first, attached by a tuft of tawny fibrils ; gills crowded, 
yellow or yellow brown, shortly adnate, attenuated outwards, narrow; 
spores white, clavate, 11-13 xg/t. Scattered or clustered, on the ground 

30 Transactions British Mycological Society 

among dead leaves and twigs. Peradeniya, no. 2408, 14 June 1907; no. 
2870, 16 July 1909. Henaratgoda, no. 5904, 25 June 1918. 

Berkeley and Broome gave the spores as subglobose, 6 p diameter. Their 
type specimen bears a few minutely warted globose spores about 
diameter, no doubt intrusive Penicillium or Aspergillus. 

Marasmius kemibaphus was described as umbonate and sulcate, but the 
sulcae are chiefly the result of drying, and the pileus is not umbonate, but 
darker in the centre than on the outer half, a fact which perhaps suggested 
the name. There was no painting of this species. Thwaites no. 204, from 
which it was described, was a mixture, part of which was Collybia leuco- 
phaea, and the painting, no. 204, is of the latter species. Marasmius chondripes, 
also part of Thwaites no. 204, is a young M. confertus. 

Marasmius haematocephalus lAont., Syll. Crypt, no. 

Pileus conico-campanulate or hemispherical, up to 7 mm. diameter, 
sometimes repand and up to 2 cm. diameter, plicato-sulcate almost to the 
centre, membranous, purple red or bluish purple or red brown; stalk hair- 
like, flexuose, usually 3-4 cm. long, 0-25-0-5 mm. diameter, horny, 
shining, black brown below, pale brown above, with a small tuft of white 
hyphae at the base; gills white, often tinged purple like the under side of 
the pileus, often with a red edge, free but close to the stalk or adnate to the 
swollen apex of the stalk, few, distant, ventricose, interstices veined wheit 
old; spores white, clavate (like an Indian club), 16-22 x 3-4 p.. On 
decaying leaves and twigs, gregarious. Common in mid and low country 
(PL IV, fig. 4). 

No. 3510, Peradeniya, 21 July 1912, was dark greenish grey or olive, but 
otherwise did not differ from the typical purple red form. 

Marasmius helvolus var. brunneolus B. & Br., Fungi of Ceylon, no. 375. 

Pileus conico-campanulate or hemispherical, about i cm. diameter, 
8 mm. high, brown in the centre, becoming yellow brown outwards, 
membranous, plicato-sulcate nearly to the centre or half-way, minutely 
velvety, the spines on the cells of the pellicle tending to be arranged in 
dots; stalk up to 4-5 cm. long, 0-5 mm. diameter, equal, glabrous, shining, 
at first pellucid, becoming brown from the base upwards, with a spreading 
tuft of white or tawny hyphae at the base; gills distant (about twelve), 
white, free, ventricose; spores white, narrow clavate, i6~ig x qp. On dead 
branches and leaves. Peradeniya, no. 2904, 8 August 1909; no. 4204, 
23 October 1914 (PI. IV, fig. 13). 

The specimens described above agree with those of Thwaites no. 752 at 
Kew and Peradeniya. Berkeley and Broome stated that the pileus and" gills 
were brownish, but there was no figure. M. helvolus Berk., type, was 
described from Cuba. 

Marasmius coniatus B. & Br., Fungi of Ceylon, no. 358. 

Pileus conico-campanulate, up to 4 mm. diameter and 3 mm. high, 
plicato-sukate, olive brown in the centre, becoming brownish buff out- 
wards, membranous, sprinkled with minute glistening particles, especially 

Revision of Ceylon Marasmii. T. Fetch 3i 

in the furrows, cells of the pellicle bearing close-set cylindrical spines up to 
6 /x high; stalk black (brown when dry), shining, hair-like, up to 2 ‘5 cm. 
long, 0-2 mm. diameter, pellucid at the apex, arising from a thin white 
layer of mycelium; gills free, distant, ascending, pallid, edge brown. On 
dead leaves and twigs. Peradeniya, no. 4221, 25 October 1914 (PL IV, 
fig. i). 

Marasmius sabconiatus Fetch, n.sp. 

Pileus up to 5 mm. diameter, broadly convex, centre umbilicate, plicato- 
sulcate to the centre, dull reddish brown, sprinkled with minute hyaline 
particles, cells of the pellicle crowned with close-set conical spines ; stalk up 
to 6 mm. long, stout, 0*4 mm. diameter, black brown, glabrous, carti- 
laginous; gills numerous (14), distant, cream-coloured, broad, lower edge 
straight or somewhat ventricose, broadly adnate. On bamboo. Pera- 
deniya, no. 4295, 12 November 1914. 

Looks like a small stout form of M, coniatus^ but differs in the number 
and attachment of the gills, and the absence of a film of mycelium on the 

Marasmius semipellucidus B. & Br., Fungi of Ceylon, no. 360. 

Pileus conico-campanulate, sometimes expanding to almost plane, some- 
times with a recurved margin, up to i cm. diameter, membranous, plicato- 
sulcate almost to the centre, pinkish brown; stalk usually about 3 cm. long, 
but sometimes up to 8*5 cm., 0*5 mm. diameter, at first light brown, 
becoming almost black at the base, horny, smooth, the base surrounded by 
a small cushion of hyphae; gills white, free, ventricose, distant (usually 12 
to 16), sometimes with a brown or reddish edge; spores white, clavate, 
14-22 X 4~5 /x. The whole pileus and gills have a pinkish tinge when moist. 
On decaying leaves and twigs. Peradeniya, nos. 2437, 2438, 16 June 1907; 
no. 4287, 9 November 1914; no. 4315, 20 November 1914 (PL IV, fig. 8). 

Berkeley and Broome described the stalk as Toto striato’, but that is not 
evident on fresh specimens. They also stated that it is distinguished by the 
upper part of the stem being pale and pellucid, a feature which is common 
to all species of this class in their earlier stage. 

Marasmius lateritius Fetch, n.sp. 

Pileus convex or conico-convex, even or obtusely umbonate, up to 6 min. 
diameter, pinkish red or brick red, darker when old, often mottled with 
yellowish patches, margin not striate or plicate (except when old) ; stalk up 
to 2*5 cm. high, i mm. diameter, at first white, then yellow brown from the 
base'upwards, horny, minutely hispid, strigose at the base; gills white, then 
yellow, not distant, rounded or truncate behind, attenuated outwards, 
free but close to the stem; spores white, clavate, 10-13 x 3-4 /x. On the 
ground among dead leaves. Peradeniya, no. 4375, 15-16 December 1914 

(PL IV, fig. 5)« 

Marasmius albocapitatus Fetch, n.sp. . ■ 

Pileus pure white, hemispherical, up to 4 mm. diameter, minutely 
umbilicate, feebly sulcate; stalk up to 1*5 cm. long, 0*2 mm. diameter, 

32 Transactions British Mycological Society 

black brown, hair-like; gills white, narrow, ascending, adnate, edge obtuse; 
spores white, narrow oval, 8— to The pileus and stalk bear long erect 

spreading hairs, not visible to the naked eye, yellow brown by transmitted 
light, acute, tapering, about 4/i diameter below, up to 0-35 mrn. long, and 
the Stalk bears in addition minute irregular thick-walled hairs, usually 
curved, up to 30 x 6/x. Cystidia on the e%e of the gills flask-shaped, up to 
34jLt high, 12 (U diameter below. On dead leaves. Hakgala, no. 4127, 
September 1914; no. 5294, April 1917 (PL IV, fig. 15). 

Marasmius micraster Fetch, n.sp. 

Pileus 3 mm. diameter, almost plane when moist, campanulate and 
radially plicate when dry, black brown to yellow brown, paler in the 
furrows, membranous, centre depressed and wrinkled, cells of the pellicle 
globose, about i6/x diameter, strongly verrucose on the outer half with 
yellow brown warts, pileus appearing minutely pruinose when magnified ; 
stalk hair-like, black brown, glabrous, shining, about i cm. long, 0-2 mm. 
diameter; gills few (about ten), white or cream, ventricose, free but close 
to the stem, edge bears flask-shaped cystidia, up to i6/n high, with a 
truncate apex; spores white, oval, one end acute, 10-12 x 4-5 /x. On 
decaying twigs. Peradeniya, no. 4195, 21 October 1914. 

Marasmius Thwaitesii B. & Br., Fungi of Ceylon, no. 383. 

Pileus in dry weather cylindrical, 2-2-5 diameter, 3 mm. high, 
dark brown, covered with spiny clusters of hairs and plicate in longi- 
tudinal ridges; when moist, broadly campanulate or almost plane, up to 
6 mm. diameter, plicato-sulcate, blackish brown in the centre, somewhat 
lighter brown along the tops of the ridges and pale brown in the furrows; 
the centre and the dark ridges retain the hair clusters for some time, but 
they ultimately disappear, first from the ridges and then from the centre ; 
the ridges are produced beyond the margin of the pileus by a spinous tuft 
of hairs; stalk up to 2-5 cm. long, 0-25 mm. diameter, rigid, dark brown, 
almost white at the apex, rough; gills about twenty, moderately distant, 
yellowish white, edge brown distally, narrow, united behind by a very 
narrow collar. On sticks and decaying palm stems. Frequent at Pera- 
deniya. No. 4353, I December 1914 (PL IV, fig. 9). 

Marasmius gordipes Sacc. & PaoL, Mycetes Malacc, p. 6, PL V, fig. 2. 

Pileus membranous, broadly convex, with or without an acute black 
umbo at the base of the umbilicus, sulcate to the centre, 3-5 mm. diameter, 
red brown; stalk filiform, very long, up to 13 centimetres long, 0-3 mm. 
diameter, somewhat rigid, striate when dry, glabrous, shining, bfackish 
brown, insititious; gills distant (up to 12), yellowish or ochraccous, 
sometimes with a reddish edge, comparatively broad, united behind into a 
collar which may fit closely round the stalk or be widely separated from it 
and shallow; spores white, narrow oval, somewhat clavatc, 8-12 X4-5/X. 
On decaying twigs and leaves on the ground. Peradeniya, no. 2902, 

9 August 1909; no. 4235, I November 1914. Not a Horsehair Blight 
(PL IV, fig. 7). 


Revision of Ceylon Marasmii. T. Fetch 

Marasmius tiibulatus Fetch, n.sp. 

Pileus cylindrico-campanulate, deeply umbilicate, radially sulcate to 
the centre, pruinose, membranous, up to 8 mm. diameter, pallid then 
brownish grey with a pale margin and a black spot at the base of the 
umbilicus; stalk up to 2 cm. long, 0-3 mm. diameter, blackish brown, 
horny, glabrous, shining, finely longitudinally striate; gills distant, white 
then pallid with a brownish grey edge, subtriangular, united behind into a 
tube, free from the stem, which descends almost to the level of the margin 
of the pileus, free edges arched ; spores white, oval or clavate, 7-9 x 3-4 /x. 
On dead leaves and twigs. Peradeniya, no. 4243, i November iq 14; also 

18 July 1909 (PI. Ill, fig. 9). 

Marasmius griseoviolaceus Fetch, n.sp. 

Pileus campanulate, up to i cm. diameter, umbilicate, plicato-sulcate, 
membranous, greyish or brownish violet or purplish brown, or almost black 
in the centre and blackish grey elsewhere, with or without a minute umbo 
at the base of the umbilicus, feebly rugose with minute black elevations 
when fresh; stalk up to 7 cm. long, 0*5 mm. diameter, attenuated upwards, 
rigid, horny, black, shining; gills white, distant (up to 14), arcuate or 
triangular, edge coloured as the pileus, united behind into a collar round 
the stem; spores white, oval, inequilateral, 8-1 1 x 6/x. Has a black repent 
rhizomorphic mycelium, 0*25 mm. diameter, and arises either from the 
rhizomorphs or from the host tissue. Peradeniya, at the base of clumps of 
Giant Bamboo [Dendrocalamus giganteus), no, 4238, i November 1914; on 
dead leaves; no. 4213, 25 October 1914 (PL IV, %. 14). 

Marasmius rotalis B. & Br., Fungi of Ceylon, no. 387; Fetch in Ann, Perad, 
VI, 50. 

Pileus 4-6 mm. diameter, hemispherical, umbilicate, generally with 
a minute umbo at the base of the umbilicus, minutely rugose, radially 
sulcate with from eight to eighteen grooves, yellow brown, greyish brown 
or ashy, membranous; stalk 2-4 cm. long, o-2“0‘5 mm. diameter, black, 
horny, filiform, shining; gills white or yellowish, distant, rather broad, 
lower edge almost horizontal, united behind into a collar round the stalk; 
spores white, narrow oval, 8--12 x 3-4 fx. On dead leaves and twgs on the 
ground, with black, glabrous, cylindric rhizomorphic mycelium about 
0*1 mm. diameter. Peradeniya and Hakgala, frequent. 

The following colour varieties have been noted. Pileus orange red, gills 
orange, stalk brown, Hakgala, no. 3725, 23 May 1913. Pileus, gills and 
Stalk orange, Hakgala, no. 4137, September 1914. Pileus greyish fawn, 
gills white, stalk black brown, Hakgala, no. 4129, 29 September 1914. 
Pileus olive brown, Hakgala, no. 4138, September 1914. Pileus white, 
umbo black and prominent, Hakgala, no. 5587, December 1917* 

Marasmius equicrinis MuelL, J, Linn, 5 'ar. xviii, 383 ; Fetch in Perad. vi, 43. 

Mycelium rhizomorphic, black, smooth, glabrous, cylindrical, about 
O’l mm. diameter, overrunning leaves and stems of living shrubs and trees 



34 Transactions British Mycological Society 

(Horse-hair Blight). Pilei arising laterally from the aerial mycelium, or 
from dead leaves and twigs on the ground. 

Pileus up to 8 mm. diameter, hemispherical, often irregularly repand, 
umbilicate, membranous, radially sulcate, yellow brown to red brown, 
with a minute black umbo; stalk 2-10 mm. long, o-i mm. diameter, 
filiform, black, glabrous; gills few (up to eight), white, then cream or 
yellowish, distant, attenuated behind, united into a collar round the apex 
of the stalk; spores white, narrow oval, inequilateral, or clavate, 10- 
14x4^.. Cells of the pellicle (as in M. rotalis) crowned with close-set, 
blunt spines. Common in- the wet low-country of Ceylon, up to an 
elevation of 1600ft. (PL III, fig. 8). 

Marasmius rigidichorda Petph, n.sp.; M. obscuratus Petch, non B. & Br., in 
Ann. Perad. vi, 56. 

Mycelium rhizomorphic, dull black, rigid, o-q-o-fi mm. diameter, over- 
running the branches of jungle shrubs. Pilei produced on the aerial 
mycelium and on the dead tissues of the host plant. 

Pileus hemispherical, umbilicate, with margin narrowly decurved, some- 
times infundibuliform, plicato-sulcate almost to the centre, coriaceous, 
dark brown in the centre, elsewhere reddish brown, becoming ashy or 
brownish white, streaked with brown fibrils and points, i cm. diameter; 
stalk up to I cm. long, i mm. diameter, blackish brown, cartilaginous, 
rough with minute fascicles of hairs ; gills distant, narrow, arched, adnate 
to a disc at the apex of the stalk, reddish, then creamy white, rather thick, 
interstices sometimes veined; spores white, narrow oval, inequilateral, 
sometimes curved at one end, 8-10 x 3-4 ja. In wet low country jungles 
from 400 to 2000 ft. elevation. Hapugastenne, nos. 2991, 3226, October 
1909. This might be known as Elephant Hair Blight. 

In Ann. Perad. vr, 56, this species was 'assigned to M. obscuratus B. & Br. 
a re-examination of the type of the latter has shown that it consists of rather 
young examples of M. proximus, and is different from the present species. 

Marasmius ascendens Petch, n.sp. 

Mycelium rhizomorphic, black, glabrous, angular, flattened and 
twisted, O'2-0-4 mm. broad, overrunning the bark of living trees up to 
a height of 40 or 50 ft. and forming a tangle among the upper branches. 
Pilei formed on fallen affected branches. 

Pileus ashy to purple grey, up to i cm. diameter, convex, then plane or 
repand, sometimes umbonate, membranous, tough, surface minutely 
rugose; stalk up to i cm. long, filiform, 0-2 mm. diameter, or flattened, 
0'4 mm. broad, glabrous, horny, black below, purple brown above; gills 
distant, adnate or very slightly decurrent, very narrow. Hakgala, nos. 
4559^ 4643^ March 1915. 

Marasmius actinophorus B. & Br., Fungi of Ceylon, no. 385; M. coronatus 
Petch, Ann. Perad. vi, 58. 

Mycelium rhizomorphic, about 0-5 mm. diameter, dark brown, clo.sely 
covered with adpressed white hairs, which are simple, up to o-6 mm. long. 

Revision of Ceylon Marasmii, T. Fetch 35 

4-6 /X diameter, septate, thick walled, overrunning shrubs and tree trunks 
up to a height of about 4 ft. Pilei produced terminally on the mycelium 
and also on dead branches or the bark of living trees. 

Pileus hemispherical, or broadly convex or almost plane, about i cm. 
diameter; centre black or dark brown, depressed, usually with a minute 
conical umbo, clothed with coarse radially arranged fibrils which project 
in. a fringe beyond the central area, elsewhere, brownish white with radial 
ridges of coarse hairs which project over the margin; hairs on the pileus, 
3 ' 5 ~ 4 :l^ diameter, smooth, simple, thick-walled, flexuose below, separate at 
the base, cohering above in pointed tapering fascicles up to 0-5 mm. long; 
stalk 4-7 cm. long, 0-25-0-5 mm. diameter, black, dull, not horny, equal, 
thickly clothed with adpressed white hairs; gills white, crowded, free, 
slightly ventricose; spores white, narrow oval, slightly inequilateral, 
7-9X4-5ja. Common at Hakgala (PL IV, fig. 16). 

Marasmius Hakgalensis Petch, n.sp. 

Pileus hemispherical, or broadly campanulate, up to 4 mm. diameter, 
purple grey or ashy brown, darker in the centre, membranous, minutely 
rugose, even, not umbilicate or umbonate, feebly striate at the margin; 
stalk up to 3 cm. long, 0-5 mm. diameter, purple brown or black, equal, 
horny, clothed with scattered, rigid, spreading, white hairs ; gills moderately 
distant, adnate, white, or pallid with a white edge. On dead leaves and 
twigs on the ground, associated with black, angular, rhizomorphic . 
mycelium, at first setulose like the stalk, becoming glabrous. Hakgala, 
common in jungle belts (PL IV, fig. 12). 

Var. denudata Petch. Stalk glabrous; gills more crowded, purpHsh 
(PL IV, fig. 12 left). 

Marasmius Paspali Petch, n.sp. 

Pileus hemispherical, not umbonate or depressed in the centre, bluish 
grey, feebly sulcate, pruinose, 2-5 mm. diameter; stalk excentric, blackish, 
white, fibrillose, cartilaginous, slightly attenuated upwards, 

2 mm. long, o-2 mm. diameter; gills ochraceous, rather broad, few (three 
or four, with short intermediates), adnate, interstices veined. On leaf- 
sheaths of Paspalum dilatatum, apparently parasitic. Old Peradeniya, no. 
6531, 12 August 1922 (PL III, fig. It). 

Marasmius Campanella Holtermann, Myk. Unters. aus den Tropen (1898), 
p. 105; ^ Cantharellus capensis" (in part) in Fungi of Ceylon, no. 350. Brown 
variety, Marasmius rufescens B. & Br,, Fungi of Ceylon, no. 3941 Xerotus 
tener B. & Br., Fungi of Ceylon, no. 426. 

Pileus circular, orbicular, or reniform, up to 4 cm. diameter, but usually 
about 2 cm., campanulate, then convex, edge often recurved, sulcate over 
the gills and veins, depressed over the stalk, glabrous, slightly rugose, 
membranous, tough, not gelatinous, white, grey or purple grey or greyish 
brown, blackening slightly when old, sometimes rufous, rarely orange; 
stalk short, e.xcentric, curved, sometimes central, 3-4 mm. long, i mm. 
diameter, solid, at first the same colour as the pileus and clothed with 

36 Transactions British Mycological Society 

minute fibrils, then black and glabrous, expanding at the apex, swollen at 
the base; gills few, usually narrow but sometimes thin and broad (up to 
3 mm.), distant, often forked, united by strong veins when old, adnate to 
the swollen apex of the stalk, the whole lower surface of the pileus slightly 
paler than the upper; spores white, oblong oval, 7-12 x 5-6 fi. On dead 
branches, stems of lianes, etc. Generally distributed and frequent (PI. II, 
fig. 14; PL III, fig. 6; PI. IV, fig. 18). 

This species was discussed in Ann. Perad. iv, 404-6 (1910), where it was 
recorded that Berkeley and Broome’s Ceylon specimens of Cmtharellus 
capensis contained at least two species, one membranous and stalked, the 
other gelatinous and usually sessile, which answered to Holtermann’s 
Marasmius Campanella and Favolaschia bispora respectively, and it was 
pointed out that Thwaites’s figure, no. 90, cited for Cantharellus capensis was 
Favolaschia bispora. The latter species was not described until 1912, when it 
appeared in Sacc. Syll. Fung, xxi, 358 as Laschia bispora (Holt.) Sacc. & 
Trav. with a spore measurement the same as Marasmius Campanella and 
certainly wrong for both. These species are said to have been collected in 
Java, but Holtermann did not give the locality, and as he worked for some 
time at Peradeniya, and the two formerly occurred in quantity near the 
entrance to the Botanic Gardens where they could scarcely have been 
overlooked by a mycologist, it would seem probable that he may have 
collected them in Ceylon. In Mycol. Mies, no. 58 (1919), 815, G. G. Lloyd 
stated that, from the figure, M. Campanella was probably in part Campanella 
cara/toa (Jungh.) Lloyd, and in no. 70 (1923), 1225, he recorded that 
Berkeley’s Ceylon determinations of Cantharellus capensis were Campanella 
cucullata. Both these statements no doubt refer to Laschia bispora, but as its 
spores are not globose it cannot ht Campanella. 

Marasmius rufo-aurantiacus Fetch, n.sp. 

Pileus at first galeate, then expanded, broadly campanulate or almost 
plane, orbicular, up to 2 cm. diameter, red brown at first, becoming 
brownish fawn or buff, smooth, thin, membranous, large specimens 
sulcate; stalk excentric, white or brownish, minutely powdered or toincn- 
tose, tomentose at the base, 2-3 mm. long, 0-3 mm. diameter; lower 
surface and gills bright orange, fading to fawn; gills adnate, arched, rather 
broad, edge thick, interstices veined, spotted with clusters of cylindrico- 
clavate cystidia crowned with red brown deposits; spores white, broadly 
clavate, slightly curved at the apex, 8-12 X4p.. On dead twigs and bark. 
Hakgala, no. 4027, April 1912; no. 4028, March 1914; no. 4130, September 
1914 (PI. II, fig. 13). 

Marasmius porphyreticus Fetch, n.sp. 

Pileus usually galeate, then almost plane, orbicular, stalk excentric, 
almost lateral, or convex, almost centrally stalked, pale or dark purple 
brown, rugose, plicato-sulcate, up to 2 cm. diameter; stalk short, i mm. 
diameter, expanding upwards, purple brown, glabrous; lower surface and 
gills pale purple brown; gills distant, adnate, about six reaching the top of 
the stalk with shorter intermediate, usually forked, edge thick; spores 

Revision of Ceylon Marasmii. T. Fetch 37 

white, narrow oval, 7-11 X3-4/X. On dead sticks. Hak^ala, no. 4126, 
28 September 1914 (PL II, fig. n). 

Marasmiiis alliariiis Fetch, n.sp. 

Pileus orbicular, up to 2 cm. broad, membranous, broadly convex, 
smooth, red brown, sulcate towards the margin when fresh; stalk excentric, 
very short, expanding upwards, longitudinally silky, white; gills cream- 
coloured, of three lengths, the longer adnate, arched, distant, broad, 
interstices veined ; spores white, oval, somewhat inequilateral, 7-1 0 x 4-5 /x. 
Strong smell of garlic when bruised. On dead wood and sticks. Hakgala, 
no. 3539 j April 1912; no. 3910, January 1914; no. 4060, April 1914; no. 
4131, September 1914; no. 5585, December 1917 (PI. II, fig. 12). 

Marasmius ignobilis B. & Br., Fungi of Ceylon, no. 390. 

Pileus orbicular, convex, pale fawn-coloured, thin, minutely rugose, up 
to 8 mm. diameter; stalk white, excentric, curved, up to 1-5 mm. long, 
0-5 mm. diameter, pruinose or subtomentose ; lower surface white; gills 
distant, adnate, edge arcuate or straight, sometimes forked, connected by 
strong veins. Old specimens ochraceous, the stalk blackening at the base. 
Strong smell of garlic when fresh. On dead tvdgs on the ground. Henarat- 
goda, no. 5346, 12 September 1917. 

No number or date was cited by Berkeley and Broome for this species, 
which was said to have occurred on dead wood in the south of the island. 
In Ann. Perad. iv, 391, I referred it to Oudemansiella subaurantiaca, basing 
that opinion on Thwaites no. 100, Peradeniya, which was cited under 
Marasmius ignobilis as a variety. There is, however, in Herb. Kew., in the 
cover of M. epochnous, Thwaites no. 683, a specimen on twigs from the 
south of the island, marked M. ignobilis, which answers to Berkeley and 
Broome’s description and that given above. 

Marasmius purpureo-albus Petch, n.sp. 

Pileus usually orbicular, up to 3 cm. diameter, stalk excentric, convex, 
then repand, sometimes centrally stalked, broadly convex, at first pale 
purple, becoming white or pale ochraceous, depressed or umbilicate over 
the stalk, irregularly plicate, glabrous; stalk white, stout, up to i cm. long, 

2 mm. diameter in the middle, equal or expanding upwards, with a ring of 
tomentum at the base; gills distant, ventricose, the longer sinuato-adnate, 
white or pallid; spores white, narrow oval, inequilateral, 10-14 x 5-6 p.. 
On a decaying banana stem, no. 5303, Peradeniya, August 1917. On bark 
scales omViVmg Lagerstroemia, no. 6541, Peradeniya, 21 September 1922; on 
fallen twigs of Lagerstroemia, no. 6557, Peradeniya, 5 November 1922 
(PL III, figs. 3, 10). 

Marasmius Amomi Petch, n.sp. 

Pileus at first convex, purple brown, tomentose with spongy tomentum 
and a few adpressed white hairs, then broadly convex, plane, or ipegularly , 
repand, up to 5 cm. diameter, irregularly radially plicate, purplish brown 
at the base, elsewhere pale ochraceous, sparsely covered with innate 

38 Transactions British Mycological Society 

radiating fascicles of purple brown fibrils, matted silky between; stalk 
excentric, white, glabrous, expanding upwards, up to i cm. long, i '5 mm. 
diameter, with a basal tuft or cushion of orange hyphae; gills white or 
pallid with a purple tinge, distant, adnate, forked, the edge irregularly 
crisped in repand specimens, interstices veined. On dead stems of 
Amomum. Hakgala, no. 4652, April 1915. 

Marasmius Calami Fetch, n.sp. 

Pileus up to 2 cm. diameter, membranous, broadly convex, strongly 
sulcate over the gills and irregularly bullate between, white with a greenish 
tinge, margin when fresh fimbriate with clavate hairs, 3 p. diameter below, 
4-8 p. above; stalk excentric, curved, connate at the base, up to i cm. long, 
I mm. diameter, equal, expanded at apex, white, longitudinally silky 
striate; gills white, rather thick, the longer distant, adnato-decurrent, often 
forked, with shorter intermediate gills, connected by strong veins; spores 
white, clavate, 13-16 x 4-5 p.. On a dead stem of Calamus. Peradeniya, 
no. 6708*, 25 December 1923. 

Marasmius pulcher (B. & Br.) Fetch, Ann. Perad. ix, 21; Cyphella pukhra 
B. & Br., Fungi of Ceylon, no. 665. 

Mycelium without anker cells, running in stout white cords over the 
stems and leaves of living shrubs; pilei produced at the edge of the cords or 
on diffuse patches of mycelium on dead leaves and bark. Pilei sessile, 
resupinate or laterally attached, sometimes shortly stalked, reniform or 
orbicular, sometimes lobed, white, becoming pale ochraceous, thin, 
tomentose, sulcate over the gills, up to 7 mm. diameter; gills radiating 
from an excentric point, few or absent on small examples, crowded and of 
medium breadth on the larger; spores white, cymbiform, 6-8 x 4 
Mycelium (Thread Blight) common in the wet low-country. No. 5269, 
from Hakgala, may be a different species ; it was found running over and 
among dead bark scales on a dead standing tree trunk and its spores were 
measured as oval or subglobose, 4-5 X gp. (PL III, fig. 4). 

Marasmius concolor P). & C. 

The Ceylon specimens attributed to this species (Thwaites no. 685) are 
Crepidotus melleus (B. & Br.) Fetch {Aschersonia mellea B. & Br.). 

Marasmius epochnous B. & Br., Fungi of Ceylon, no. 393; ‘A/, epochnous 
B. & C.’ (in error) . 

This species was said to be white. The pilei in the type are gregarious, up 
to 3 mm. diameter, orbicular, convex, tomentose; stalk excentric, almost 
lateral, curved, up to 0-5 mm. long, 0-15 mm. diameter, each arising From 
a circular patch of radiafing hyphae; gills broad, adnate, distant. The 
pileus and gills are very friable, and it does not seem a good Marasmius. It 
was collected in the south of the island and has not been matched by recent 

Revision of Ceylon Marasmii. T. Fetch 39 

specimens. It probably grew on the under surface of a branch as^ in the' 
herbarium specimen, the stalks curve towards the substratum, so that the 
lower surface of the pileus is uppermost. 

Marasmiiis {Mycena) galericula Ces. 

This species was collected by Beccari in the Botanic Gardens, Pera- 
deniya, and was described by Cesati as ‘^subfuniculate, lignicolous; stalk 
three inches long, filiform, horny, rigid, brown, flexuose (not twisted), 
smooth, scarcely pulverulent or pruinose, but opaque; pileus campanulate, 
one inch diameter, colour when dry lurid, with the appearance of an 
agaric of the series Mycenae rigidipes^ collapsed when old. From the de- 
velopment of the stalk I am doubtful as regards MarasmiusJ' Mycenae 
rigidipes is Fries’s section of Mycena which includes Mycena galericulata^ and 
from Cesati’s choice of a name it might be surmised that he saw some 
resemblance to that species. I am unable to suggest what it may have been. 

Marasmius inustus B. & Br., Fungi of Ceylon, no. 384. 

The type resembles rather small specimens of M, Campanella^ somewhat 
thicker than usual, and stained black here and there, as though it changed 
colour when bruised. They have not been matched by fresh specimens and 
may be a distinct species. They are not gelatinous. 

Marasmius prasimis B. & Br., Fungi of Ceylon, no. 373. 

Berkeley and Broome described this as pileus 3 cm. across, of a delicate 
greyish green, somewhat fleshy, glabrous, margin irregular, sulcate, flesh 
white; stalk 3 cm. long, 1-5 mm. thick; gills narrow, arcuate, decurrent, 
ochraceous; on dead twigs, etc. No Thwaites number, locality or date was 
cited, and there was no painting. The type in Herb. Kew. consists of a 
single pileus pressed flat, about 4 cm. diameter, extremely thin. It bears 
on the lower surface radial bands of loose tissue, bounded by denser lines, 
and examination shows that the gills have been cut away, leaving the 
trama bounded by their bases. Pieces of the gill remain towards the margin, 
but these do not show anything which would afford a clue to its identity. 
It would seem that the specimen has been cut up for preservation, but if so 
the other pieces are not in Herb. Kew. It is not possible to determine 
anything from the specimen, and I never found anything to fit the de- 

Marasmius proletarius & C. 

This species from Cuba was described as white. The Ceylon painting, 
Thwaites No. 1 1 70, shows a coloured pileus and represents young specimens 
of 3 /. coniatus. The name should be deleted from the Ceylon list. 

Marasmius radians B. & Br., Fungi of Ceylon, no. 366. 

Thwaites no. 103 was divided by Berkeley and Broome into two species, 
Cantharellus inaequalis B. & Br., Fungi of Ceylon, no. 347, and Marasmius 
radians B. & Br. There was no painting. The two appear to be the same, 
and a Cantharellus, but until fresh specimens have been collected, the 
question may remain open. Both grew on wood or sticks. 

40 Transactions British Mycological Society 

Berk., Decades of Fungi, no. 156. 

Pileus broadly convex, depressed, ochraceous, thin (? through drying), 
about 2 cm. diameter, minutely scurfy, no specia.1 pellicle; stalk com- 
pressed, cartilaginous, minutely pruinose, white, strigose at the base, with 
tawny mycelium spreading over the substratum, about 2 cm. long, stout 
when fresh, i mm. broad in the dry specimen; gills ventricose, adnate, 
moderately crowded; spores yellow brown, oval or obliquely oval, 
apiculate, 6 x 4 /x,. On rotten wood. 

This species has not been identified again. It was sent by Gardner, with 
a figure which shows a stout white stalk and a depressed ochraceous pileus. 
The radial grooves noted by Berkeley are probably the result of drying. 
It appears to be a Naucoria. 

Marasmius Wynniae var. auroricolor B. & Br., Fungi of Ceylon, no. 353- 

This is a Mycena, which may be known' as Mycena auroricolor Fetch, 

Solitary or caespitose; pileus up to 2-5 cm. diameter, broadly convex, 
then almost plane, slightly depressed in the centre, margin striate when 
moist, pale lilac, rose in the centre, becoming entirely rose pink, or when 
young everywhere rose pink with a purplish tinge, pallid when old, 
sprinkled with minute glistening particles when dry; flesh thin, white in the 
centre; gills paler than the pileus, adnate or adnato-decurrent, narrow, 
3 mm. broad, attenuated outwards, lower edge straight, interstices 
strongly veined; stalk up to 5 cm. long, 2-5 mm. diameter, equal, coloured 
like the pileus, hollow, smooth, shining, brittle, sometimes ridged above 
with decurrent lines from the gills, slightly strigose at the base; spoz-es 
white, oval, 6-8 x 4 jti. On the ground among vegetable refuse. Peradeniya, 
I July 1908; I August 1909; also no. 4142. 

The specimen Thwaites no. 204 cited by Berkeley and Broome as ‘ the 
same thing’ is M. leucophaeus. 

Latin descriptions of new species 

Marasmius purpureo-griseus Fetch, sp.nov. Fasciculatus ; pileo late convexo, 
margine pallidiori, minute radiatim sericeo, ca. 1-5 cm. diam., margine 
incurvato; carneo albo tenui; stipite ad 4 cm. alt., i -5 mm. diam., 
brunneo-albo, nitenti, cartilagineo, farcto dein cavo, fistuloso; lamcllis 
confertis, albis vel cremeis, latis, adnatis v. sinuato-adnatis, acie recta. 
Ad lignum emortuum, mycelio albo affixus. Henaratgoda, Ceylon. 

Marasmius rufo-ochraceus Fetch, sp.nov. Fasciculatus v. sparsus ; pilciz late 
convexo v. fere piano et undulato, ad 3-5 cm. diam., centro rufo-brunneo, 
ad marginem ochraceo v. pallido, sicco toto ochraceo, radiatim sulcato, 
glabro, tenui, cartilagineo; stipite ad 3 cm. alt'., compresso, ad 3 mm. hit., 
cartilagineo, rufo-bruimeo, supra pallidiori, glabro v. minute tomentoso, 
farcto dein cavo; lamellis subremotis, subventricosis, adnatis, pallidisj 
bruimescentibus, aetate interstitiis valde venosis; sporis albis, clavatisj 
10-14 X 5-6 p. Ad lignum emortuum. Hakgala, Ceylon. 

Marasmius brumeostriatus Fetch, sp.nov. Pileo campanulato dein fere 
piano, obtuse umbonato, ad 2 cm. diam., rufo, centro saturation, margine 

Revision of Ceylon Marasmii. J. Fetch 41 

pallidiori flavo-bmnneo, radiatim sulcato, sulcis saturatioribus, minute 
rugoso, margine saepe repando; stipite ad 2 cm. alt., i mm. diam., apice 
incrassato, cartilagineo, rufo-brunneo, pruinoso v. minute tomentoso, 
basi tomentoso, e strato tenui albo oriundo ; lamellis pallido-ochraceis, 
distantibus, crassiusculis, angustis, aequalibus, adnatis ; sporis albis, clavatis 
V. ovalibus, 5-9 x 3-4 /x. Ad folia et ramulos dejectos. Hakgala, Ceylon. 

Marasmius senescens Fetch, sp.nov. Pileo i -5 cm. diam., late convexo, centro 
depresso, interdum umbonato, plicato-sulcato, centro corrugato, sordide 
griseo margine pallidiori, tenui, centro lender tomentoso, alibi sparsim 
fibrilloso et hinc illic consperso; stipite i cm. alt., 1-5 mm. diam., 'supra 
expanso, albo v. cinereo, fuscescenti, compresso, striato, minute longi- 
tudinaliter sericeo, cartilagineo; lamellis distantibus, adnatis v. adnato- 
decurrentibusy circum apicem stipitis connexis et separantibus, latis, 
leniter ventricosis, crassiusculis, furcatis, pallidis, brunnescentibus, inter- 
stitiis venosis; sporis albis, angusto-ovalibus, subpyriformibus, 7-10 x 3-4 /x. 
Ad terram. Peradeniya, Ceylon. 

Marasmius stypinoides Fetch, sp.nov. Pileo albo, centro cinereo, late con- 
vexo V. fere piano, ad 5 mm. diam., umbilicato, leniter radiatim sulcato, 
minute rugoso, margine crenato; stipite ad 8 mm. alt., 0-4 mm. diam., 
saepius curvato, primo albo dein nigrescenti, pulverulento, striato, apice 
incrassato; lamellis albis, confertis, subventricosis, acie serrato. Ad 
corticem emortuum. Hakgala, Ceylon. 

Marasmius cineraceus Fetch, sp.nov. Pileo infundibuliformi, margine 
decurvo, leniter radiatim sulcato, 9 mm. diam., cinereo, levi, pellicula 
hypharum intertextarum; stipite nigro, albo-pruinoso, cartilagineo, 
I cm. alt., 0*75 mm. diam., aequali, e disco glabro rotundato oriundo; 
lamellis subconfertis, albis, sigmoideis, latis, adnato-decurrentibus, acie 
setrato, interstitiis venosis. Ad folia emortua. Peradeniya, Ceylon. 

Marasmius subconiatus Fetch, sp.nov. Pileo ad 5 mm. diam., late convexo, 
plicato-sulcato, umbilicato, obscure rufo-brunneo, particulis hyalinis 
consperso, cellulis pelliculae spinis conicis confertis coronatis ; stipite ad 
6 mm. alt., crasso, 0*4 mm. diam., nigro-brunneo, glabro, cartilagineo; 
lamellis numerosis (14), distantibus, cremeis, latis, acie recta v. subventri- 
Gosis, late adnatis. Ad bambusam emortuam. Peradeniya, Ceylon. 

Marasmius lateritius Petch, sp.nov. Pileo convexo v. conico-convexo, piano 
V. obtuse umbonato, ad 6 mm. diam., pallide rubra v. lateritio, saepe 
maculis flavis; stipite ad 2-5 cm. alt., i mm. diam., primo albo dein flavo- 
brunneo, corneo, minute hispido, basi strigoso; lamellis albis dein flavis, 
non distantibus, postice rotundatis v. truncatis, liberis sed stipitem 
approximatis ; sporis albis, clavatis, 10-13 Ad terram inter folia 

emortua. Peradeniya, Ceylon. 

Marasmius albocapitatus Petch, sp.nov. Pileo candido, hemisphaerico, ad 
4 mm. diam., minute umbilicato, leniter sulcato; stipite i *5 cm. alt, 
0'2 mm. diam., nigrobrunneo, i^etaceo; lamellis albis, angustis, adscenti- 
bus, adnatis, acie obtuso; sporis albis, anguste ovalibus, B-iO x 3 jtx; pileo et 
stipite crines longos patentes flavo-brunneos, attenuatos, acutos, infra 4fx 
diam., ad 0*35 mm. alt. ferentibus; stipite etiam crinibus niinutis, irregu- 
laribus, saepius curvatis, ad 30 x 6/x, vestitis; cystidiis acie lamellarum 

42 Transactions British Mycological Society 

ampullaceis, ad 34 /tt alt., infra 12 diam. Ad folia emortiia. Hakgala, 

Marasmius micr aster Fetch, sp.nov. Pileo 3 mm. diam., udo fere piano, 
sicco campanulato et radiatim plicate, nigrobrunneo v. flavobrunneo, in 
sulcis saturation, membranaceo, centre depresso et corrugate, cellulis 
pelliculae ca. 16/x, diam., externe valde verrucosis; stipite setaceo, nigro- 
brunneo, glabro, nitenti, ca. i cm. alt., o-2 mm. diam.; lamellis paucis, 
albis V, cremeis, ventricosis, liberis; cystidiis ampullaceis, ad 16 /i- alt,, apice 
truncatis; sporis albis, ovalibus, uno fine acutis, 10-12 x ramulos 

emortuos. Peradeniya, Ceylon. 

Marasmius tubulatusFttchy sp.nov. Pileo cylindrico-campanulato, profiinde 
umbilicato, plicato-sulcato, pruinoso, membranaceo, ad 8 mm. diam., 
pallido dein brunneo-cinereo, margine pallidiori, centre nigro; stipite ad 
2 cm. alt., 0*3 mm. diam., nigro-brunneo, corneo, glabro, nitenti, longi- 
tudinaliter striato ; lamellis distantibus, albis dein pallidis, acie brunneo- 
griseo, subtriangularibus, postice in tubulo junctis; sporis albis, ovalibus 
V. clavatis, 7-9 x 3-4 /x. Ad folia et ramulos emortuos. Peradeniya, Ceylon, 

Marasmius griseo-violaceus Fetch, sp . no v. Pileo campanulato , ad i cm , diam . , 
umbilicato, plicato-sulcato, membranaceo, griseo-violaceo v. purpureo- 
brunneo v. centre fere nigro alibi nigro-griseo, udo minute nigro-rugoso ; 
stipite ad 7 cm. alt., 0*5 mm. diam., supra attenuate, rigido, corneo, nigro, 
nitenti; lamellis albis, distantibus, arcuatis v. triangularibus, acie colorati, 
in collarium liberum postice conjunctis; sporis albis, ovalibus, inaeqiii- 
lateralibus, 8-1 1 x6 /x. Mycelio nigro rhizomorphoideo, 0-25 mm. diam, 
Dendrocalami gigantei et ad folia emortua. Peradeniya, Ceylon. 

Marasmius rigidichorda Fetch, sp.nov. Pileo hemisphaerico, umbilicato, 
margine anguste decurvo, interdum infundibuliformi, plicato-sulcato, 
coriaceo, centre fusco-brunneo, alibi rubro-brunneo, fibris et punctis 
brunneis radiato, 1 cm. diam.; stipite ad i cm. alt,, i mm. diam., nigro- 
brunneo, cartilagineo, scabro; lamellis distantibus, angustis, arcuatis, 
adnatis, rubescentibus dein cremeis, crassiusculis; sporis albis, angusto- 
ovalibus, inaequilateralibus, 8-10 x 3-4 Mycelio aerio rhizomorphoideo, 
nigro, rigido, o*4-0‘6 mm. diam. Ad frutices. Hapugastenne, Ceylon, 

Marasmius ascendens Fetch, sp.nov. Pileo cinereo v. purpureo-griseo, ad 
I cm. diam., convexo dein piano v. repando, interdum umbonato, 
membranaceo, tenaci, minute rugoso; stipite ad i cm. alt., 0*2 mm. diam,, 
setaceo, v. compresso, 0-4 mm. lat., glabro, corneo, infra nigro, supra 
purpureo-brunneo; lamellis distantibus, adnatis v. leniter decurrentibus, 
angustissimis. Mycelio aerio rhizomorphoideo, nigro, glabro, angulato, 
compresso et torto. Ad truncos et ramos arborum vivorum. Hakgala, 

Marasmius Hakgalensis Fetch, sp.nov. Pileo hemisphaerico v. late campa- 
nulato, ad 4 mm. diam.; purpureo-griseo v. cinereo-brunneo, centre 
saturatiori, membranaceo, minute rugoso, margine leniter striato; stipite 
ad 3 cm. alt., 0*5 mm. diam., purpureo-brunneo v. nigro, aequali, corneo, 
setuloso; lamellis subdistantibus, adnatis, albis, v. pallidis acie albo. 
Mycelio rhizomorphoideo, terrestri, nigro, angulato, setuloso dein glabro. 
Ad folia et ramulos emortuos. Hakgala, Ceylon. 

Revision of Ceylon Marasmii, T. Patch 43 

Var. demdata Fetch. Stipite glabro; lamellis magis confertis, pur- 

Marasmius Paspali^ Fetch, sp.nov. Fileo hemisphaerico, coeruleo-griseo, 
leniter sulcato, pruinoso, 2*5 mm. diam.; stipite excexitrico, nigrescenti, 
fibrilloso, cartilagineo, 2 mm* alt., 0-2 mm, diam. ; lamellis ochraceis, 
latiusculis, paucis, adnatis, interstitiis venosis. Ad vaginas Paspali dilatati. 
Feradeniya, Ceylon. 

Marasmius rufo-aurantiacus Fetch, sp.nov. Fileo primo galeato, dein 
expanse, late campanulato v. fere piano, orbicular!, ad 2 cm. diam., 
primo^ rubro-brunneo, dein brunneo-cervino, levi, tenui, membranaceo, 
majoribus sulcatis; stipite excentrico, albo v. brurmescenti, minute 
pulverulento v. tomentoso, basi tomentoso, 2-3 mm. alt,, 0*3 mm. diam.; 
lamellis vivide aurantiacis dein cervinis, adnatis, arcuatis, latiusculis, acie 
crasso, interstitiis venosis, acervulis cystidiorumcylindrico-clavatorummassis 
rufis coronatorum maculatis ; sporis albis, late clavatis, apice leniter curvatis, 
8-12 X Ad corticem et ramulos emortuos. Hakgala, Ceylon. 

Marasmius ppTphyreticus Fetch, sp.nov. Fileo saepius galeato, dein fere piano, 
orbiculari, stipite excentrico, v. convexo, stipite fere centrali, purpureo- 
brunneo, rugoso, plicato-sulcato, ad 2 cm. diam.; stipite brevi, i mm. 
diam., supra incrassato, purpureo-brunneo, glabro; lamellis paucis, 
purpureo-brunneis, distantibus, adnatis, saepius furcatis, crassis; sporis 
albis angusto-ovalibus, 7-11 x 3-4 p,. Ad ramulos emortuos. Hakgala, 

Marasmius alliarius Fetch, sp.nov. Fileo orbiculari, ad 2 cm. lat., late 
convexo, membranaceo, levi, rufo-brunneo, margine sulcato; stipite 
excentrico, brevissimo, supra incrassato, longitudinaliter sericeo, albo; 
lamellis cremeis, majoribus adnatis, arcuatis, latis, interstitiis venosis; 
sporis albis, ovalibus, 7-10 x 5 ft. Contuso alliatus. Ad lignum et' ramulos 
emortuos. Hakgala, Ceylon. 

Marasmius purpureo-albus Fetch, sp.nov, Fileo saepius orbiculari, ad 3 cm. 
diam,, convexo, repando, stipite excentrico, interdum stipite centrali, late 
convexo, primo pallide purpureo, dein albo v. pallide ochraceo, postice 
depress© v. umbilicato, vage plicato, glabro; stipite albo, crasso, ad i cm. 
alt., medio 2 mm. diam., aequali v. supra incrassato, basi annulari- 
tomentoso; lamellis distantibus, ventricosis, majoribus sinuato-adnatis, 
albis V. pallidis; sporis albis, angusto-ovalibus, inaequilateralibus, lo- 
14x 5-6 ft. Ad caulem marcidum Musae (Banana), et ad corticem vivae 
Lagerstroeiniae. Feradeniya, Ceylon. 

Aiarasmius AmomiPtldk.^-^.noy. Fileo primo convexo, purpureo-brunneo, 
spongiGSo-tomentoso, dein late convexo v. piano v. vage repando, (ad 
5 cm. diam., vage radiatim plicato, postice purpureo-brunneo, alibi pallide 
ochraceo, innatis radiantibus fasciculis purpureo-brunnearum hypharum 
sparsim induto; stipite excentrico, albo, glabro, supra incrassato, ad i cm. 
alt., I *5 mm. diam., basi aurantiaco-tomentoso; lamellis albis v. pur- 
purascentibus, distantibus, adnatis, furcatis, interstitiis venosis. Ad caules 
emortuos Hakgala, Ceylon. 

Marasmius Calami Fetch, sp.nov. Fileo ad 2 cm. diam., late convexo, 
membranaceo, super lamellis valde sulcato, inter vage bullato, viridi-albo. 

44 Transactions British Mycological Society 

margine udo fimbriato crinibus davatis infra 3 ft diam., supra 4-8 /x; 
stipite excentrico, curvato, connexo, ad i cm. alt., i mm. diam., aequali, 
apice incrassato, sericeo-striato ; lamellis albis, crassiusculis, distantibus, 
adnato-decurrentibus, saepe furcatis, venis validis connexis; sporis albis, 
davatis, 1 3-1 6 x 4-5 /x. Ad caulem emortuum Calami. Peradeniya, 


(All figures natural size, unless otherwise stated; numbers in brackets are those of the specimens 
from which the figures were drawn.) 

Plate II 

Fig. I . M. nephelodes^ and section. 

Fig. 2. M, rivulosus, full-grown, young, and section (4380). 

Fig. 3. M. Caryotae, and section. 

Fig. 4, AT. and surface view of pileus (4294). 

Fig. 5. Af. and section (4382). 

Fig. 6. M. crispatiis, and under-surface (4239). 

Fig. 7. M. brumeostriatus, and section (4116). 

Fig. 8. AT. and section (41 15). 

Fig. 9. M. proximus (4152). 

Fig. lo. M, corticigena, full grown, section, and young (4370). 

Fig. II. M, porphyreticus, upper and under-surface (4126). 

Fig. 12. Af. upper and under-surface (4131). 

Fig. 13. AT. mfo-aurantiacusy under and upper surface (4130). 

Fig. 14. Af. Campanella, small specimen (4124). 

Plate III 

Fig. I. AT. congregatus, and section. 

Fig. 2. M, leucophaeusy large example (4215). 

Fig. 3. M. purpureo-albus, upper and under-surface, from banana (ci202). 

Fig. 4. M. pulchery X2 (5269). 

Fig. 5. AT. arboreal form, under-surface, x 2 (6542). 

Fig. 6. Af. Campanellay ochraceous form (4748) . 


Fig. 8. Af. equicrinisy and under-surface. 

Fig. 9. M. tubulatus 

Fig. 10. M. purpureO’-albuSy CtxiXxdX (6557) and lateral (6541) stalked, from Lagerstroemza. 

Fig. II. M.Paspahy X2 (6531). 

Fig. 12. AT. m'yajzty (6552). 

Fig. 13. M. hirtellus var., and upper surface, x 2 (5807). 

Fig. 14. Af. subcinereus (6627). 

Fig. 15. Af. and section (6256). 

Fig. I . 
Fig. 2. 


Fig. 4. 
^’ig- 5- 
Fig. 6. 
Fig. 7. 
Fig. 8. 
Fig- 9- 
Fig. 10. 
Fig. u . 
Fig. 12. 
Fig. 13, 
Fig. 14. 
Fig. 15. 
Fig. 1 6. 
Fig. 17. 
Fig- 18. 

Plate IV 

M. coniatus 

Af. LeveillianuSy and section (4625). 

M. Jiorideus, lyge example (4207) , and section of small example (428 1) . 

Af. haematocephaltis (4102). 

Af. and section (4375). 

M.fulviceps Uoyy). 

Af. gordipesy and under-surface (4235) . 

M, semipellucidusy and section (4305) . 

Af. and under-surface (4353). 

Af. AyAc»cAro2<f(?j, large example (4244). 

Af. small example and section, 

(right) (4133), and var. dfina&ta (left) (412'i). 
M.helvolmY 2 xo.brunneolus{d^ 204 ). 

Af. griseoviolaceusy and under-surface (4238). 

Af. and under-surface (4127). 

M. actinophorus {/^i^/^, 

M andsection (right) of young example (4248). 

{Accepted for publication iQ October 1946) 

Trans. Brit. Myc. 'Soc, 

A, George Alwis del. 

Ceylon Marasmii 

Trans. Brit. Myc. Soc 

A. George Alwis del, 

C4eylon Marasmii 

[ 45 ] 


By a. Seale-Hqyne Agricultural College^ Newton Abbot 

[Now at National Agricultural Advisory Service^ Quarry Dene^ Leeds 6) 

The Dutch rules for forecasting blight epidemics 

Many observations have been made on the relation between weather 
conditions and the dates of outbreak of potato blight epidemics. Very 
little of practical value was discovered until van Everdingen (1926)5 from 
an examination of Dutch weather data, established four conditions which 
must be satisfied before outbreaks of blight occur. The value of this work 
was that, for the first time, it enabled forecasts to be made of the probable 
occurrence of blight and, therefore, of the date when spraying to control 
the disease should be begun. 

Van Everdingen’s conditions (which will be referred to here as the 
Dutch rules) were as follows : 

(1) Dew occurring for at least four hours during the night. 

(2) Minimum night temperature not less than 10® C. (50° F.). 

(3) Mean cloudiness of the day following the dew not less than 0*8. 

(4) Rainfall on the day following the dew of at least o*i mm. 

If any day occurred on which all these conditions were obeyed, blight 
usually made its appearance within a fortnight. 

These rules have been applied successfully in Holland and a spray 
warning service organized there (van Poeteren, 1928, 1931 ; van Everdingen 
1935). They have also been tested in other countries. Wilfshire (1931) 
applied them to data from a number of places in England and Wales for 
the years 192 1-7. Although the rules were often obeyed, the results on the 
whole were less satisfactory than in Holland. Better results were obtained 
when slight deviations were permitted in the application of the rules (e.g. 
three hours’ dew formation instead of four, or 49"^ F. instead of 50° F.), 
and such a procedure seemed reasonable because the conditions governing 
biological phenomena can rarely be expressed with rigid arithmetical 
precision. This has the practical drawback, however, that it becomes much 
more difficult to apply a spray warning service with confidence because 
there is a large increase in the number of almost favourable days which are 
not followed by blight within a fortnight. I have applied the Dutch rules 
to the weather data at Newton Abbot, Devon (Beaumont, r929”32) and, 
as will be seen, have come to the same conclusion as Wiltshire. The Dutch 
rules were satisfactorily applied by Dufrenoy (1933) at Bordeaux in 1932 
and by Naoumova (1935) in the neighbourhood of Leningrad in 1933 and 

It may be useful at this stage to discuss the phytopathological significance 
of the rules. Thanks to the work of Melhus (1915), Crosier (1934) and 
others, we have a great deal of information about the biology of the blight 

46 Transactions British Mycological Society 

fungus which is directly relevant to the rules. Blight appears in the crop 
as a result of planting one or more infected tubers, in the aerial shoots from 
which the blight fungus develops as soon as conditions are favourable. 
There are two stages preceding the first visible appearance of blight in 
what may be called primary infection foci, namely, growth of mycelium in 
the shoot and formation of conidia. The first stage is governed only by 
temperature, which is covered by the second of van Everdingen’s rules, and 
which will have a cumulative effect during the growth of the plant. 
Crosier (1934) has shown that the fungus grows most rapidly at 18-21° C. 
(64-69° F.) and very slowly below 9° C. (48° F.) or above 24° C. (75° F.). 
The formation of conidia is governed both by temperature and humidity : 
the optimum temperature is i8-2Q° C. (64-72° F.). 

The other three rules of van Everdingen relate to the dissemination of 
blight rather than to the formation of primary infection foci. Naturally, if 
conditions were quite unfavourable to the spread of blight, the disease 
would not be reported, as it is unlikely that during ordinary field observa- 
tion the primary infections would be noticed. It is well known that in- 
fection by Phjtophthora infestam takes place most readily when zoospores are 
formed, and this occurs only when there is a film of moisture on the leaf. 
The longer the film of moisture persists the greater is the chance of in- 
fection. Dew formation (rule i) followed by cloudy skies (rule 3) and a 
certain amount of rain (rule 4) mean that the moisture will last a con- 
siderable time and the rules therefore allow for the conditions under which 
blight will spread sufficiently to form a noticeable outbreak. 

Application of the Dutch rules to South Devon 

Difficulties arise when using the Dutch rules owing to the nature of the 
meteorological data to be consulted. Dew formation cannot be directly 
observed, but the period may be estimated by comparing the continuous 
temperature record with the dew points recorded at 21 hr. (the night 
before) and at 9 hr. in the morning. It is by no means certain how closely 
the period obtained in this way corresponds with the actual period of dew 
formation on potato leaves. Dew does not readily form on potato leaves; 
much less readily, for example, than on grass. At most stations continuous 
temperature records are not available and in Table i dew formation has 
been regarded as occurring if the minimum temperature (read at 9 a m.) 
was more than 2° F. below the dew point (also read at 9 a.m.). 

The rneteorological data for the amount of cloud usually refer to one 
observation only (9 a.m.) and in no case to more than three, and in 1931 
they ceased to be recorded at most English stations. The sunshine record 
would seem to give a better idea of the general cloudiness during the day 
and in Table i a maximum of five hours’ sunshine has been allowed. 

The rainfall condition is most variable in its application. A combination 
of a little rain and little sunshine generally means persistent moisture on the 
leaves. But rain in the morning is often succeeded by wind and inter- 
mittent sunshine during the afternoon, and potato leaves dry up very 
readily under these conditions. It cannot be expected that the variety of 






1 . 


Weather and Outbreak of Potato Blight, A. Beaumont 47 

weather conditions can adequately be summed up in a simple rule^ and 
these limitations must be accepted. 

Another difficulty is that the nearest meteorological station for which 
recorded data are available may be some distance from the potato field. 
This was unavoidably so in Wiltshire’s observations, but in the Newton 
Abbot results given below the meteorological instruments were either in or 
very close to the potato field. 

In arriving at Table i the Dutch rules have therefore been modified as 
follows : . 

(1) Minimum temperature not less than 2° F. below the dew point. 

(2) Minimum temperature not less than 50^^ F. 

(3) Sunshine of the following day not more than five hours. 

(4) Rainfall of the following day at least o-i mm. 

Table i. 

South Devori 











1 1 July 








2 Aug. 








10 June 








15 July 







29 July 








16 Aug. 








26 Aug. 







25 July 








20 July 








8 Aug. 








24 July 







Mean 42*5 







Mean 31 


la, {The 


excluding 1931 and excluding May 

and June) 










































— ^ 







' — 






' : J ■ 







; ■"■'/22 ■ 



























Mean 22 



■'■'7' . 87 , 

Mean 12 

Column 2. Date of outbreak of blight. 

3. No, of critical periods preceding outbreak (Dutch rules). 

4. No. of critical periods in the fifteen days preceding outbreak (Dutch rules) ^ 

5. No. of days between first critical period and outbreak, excluding May (Dutch 


6. 7 and 8. As 3, 4 and 5 respectively but determined according to the temperature- 

humidity rule. 

The data obtained from the weather records (not published here owing 
to lack of space) are summarized in Table i. Examination of columns 3, 

48 Transactions British Mycological Society 

4 and 5, obtained by the application of the Dutch rules to the weather 
data, shows that the outbreak of blight was preceded within fifteen days by 
one or more critical days in every year from 1929 to 1 939. If there had been 
only one such day before the blight outbreak an accurate forecast of the 
latter could have been made. With two or three critical days within the 
fifteen-day period, it would still be possible to give a useful and closely 
approximate forecast. But in every season critical days occurred which 
were more than fifteen days before the outbreak and in every year except 
1931 the first critical day recorded was, in fact, a much longer period ahead 
of the blight outbreak. There were indeed forty-four such days out of a total 
of seventy-one, or 62 %. Even if we consider a twenty-one-day interval as of 
practical value (i.e. forecasts of blight would take the form of possible out- 
breaks any time up to three weeks from the date of forecast) the percentage 
of premature forecasts would still be as much as 56 %. 

A possible explanation of the number of premature forecasts is that the 
chances of blight appearing in epidemic form vary in the different months 
according to the growth of the crop. Obviously the chances of blight 
appearing before the shoots are through the ground are nil, whatever the 
weather, but as growth proceeds and leaves and shoots become more 
numerous the chances of blight will become greater. In other words, a 
favourable period in July is more likely to be followed by blight than one in 
June, while in May the chances are still less. Critical days are rare in May 
because the temperature is generally too low (only one day in each of seven 
seasons out of the eleven). In the seasons under investigation blight 
appeared in June only once (1931). In Table i a critical days earlier than 
July are disregarded. We now find that all the critical days are followed by 
blight within fifteen days in three seasons out of ten, and within twenty-one 
days in six seasons. The percentages of premature forecasts are reduced to 
37 and 15% respectively. This is a big improvement. Apart from 1931, 
therefore, good results were obtained when considering only the July and 
August data. It is, however, important to be able to forecast when blight 
may come exceptionally early, as it did in 1931, because spraying earlier 
than usual would Ixave been essential in that year. The application of the 
Dutch rules does not indicate why 1931 was so exceptional. 


The Dutch rules do not take into account the relative humidity of the 
atmosphere, perhaps because adequate data were not available. The 
importance of high humidity in relation to the formation of comdia is well 
known and should be considered (Beaumont, 1930). This was done, and it 
was found that humidity and temperature alone were sufficient to give a 
practically useful indication of the likelihood of blight. When the data are 
available the application of a 'temperature-humidity rule' is much 
simpler than the use of the Dutch rules. 

When this rule was first adopted relative humidity records were being 
made at 9, 15 and 21 hr. and these enabled a fairly reliable picture to be 
obtained of the degree of humidity throughout the day. Unfortunately 

Weather and Outbreak of Potato Blight. A. Beaumont 49 

only the 9 hr. reading was made from 1936 onwards, and as relative 
humidity usually diminishes during the afternoon such restricted data are 
of no value for our present purpose. This restricts the application of new 
rules to the few stations that possess continuous recording instruments or 
take sufficient hygrometric readings during the day. It is true that the 
absolute humidity of the air varies little during the day, and the relative 
humidity, which depends on temperature, can be calculated if the latter is 
known. Nevertheless, it would be a great help if direct readings of relative 
humidity at 15 hr. could be re-started. The Meteorological Office (1928) 
recognized that important variations in relative humidity occur during 
the hours of daylight between 7 and 21 hr. and are not adequately 
disclosed by observations at 7, 8, 9, 18 or 21 hr. An observation in the 
early afternoon is required to indicate the extent of the fluctuations’. The 
13 hr. reading given in the Daily Weather Map goes some way to supply 
this need. 

During a period of high humidity there is an absence of wind, and the 
drop in temperature that always occurs at night is often sufficient to induce 
dew formation, while the film produced does not dry up readily owing to 
the dampness of the air. This is especially likely when the high humidity 
persists for forty-eight hours. In effect, therefore, the humidity conditions 
comprise in one rule the Dutch rules 1, 3 and 4. 

The new conditions proposed may be stated as 

(1) Minimum temperature not less than 50° F. 

(2) Relative humidity not below 75 % for at least two days. 

Table i gives the number of critical periods preceding the outbreak of 
blight in each year, a critical period being defined as one in which the 
required temperature and humidity were both observed for two or more 
consecutive days. For comparison with the Dutch rules the total number of 
periods and the number of periods up to fifteen days before the outbreak 
are both given, as well as the number of days after the first critical period 
when blight was noted. In 1929 and 1930 only three humidity readings, at 
9 a.m., 3 p.m. and 9 p.m., were available, but in subsequent seasons the 
data were taken from ' a recording hygrometer kept in a screen placed 
among the potatoes. 

Columns 6, 7 and 8 show that blight was preceded by one or more 
critical periods, not in every season, as with the Dutch rules, but in nine 
seasons out of eleven. On the basis of the temperature-humidity rule, 
therefore, no spray forecast would have been issued in 1934 and 1937. 
Actually the epidemics were very slight in those years, so that the practical 
disadvantages of the failure to forecast are not great, a point that will be 
discussed further below. The temperature-humidity rule has the advantage 
over the Dutch rules in that in any given season the number of critical 
periods is smaller, so that a forecast can be made with more certainty 
(compare column 6 with column 3) . The table shows that in 1929 and 1938 
there was only one critical period, and in 1931 all the three critical periods 
were within the fifteen days’ period. In these three seasons, therefore, 
unequivocal forecasts could have been issued. In 1939 the single critical 
period was within the twenty-one-day interval. In four seasons out of 



50 Transactions British Mycological Society 

eleven, therefore, blight followed all the critical periods within twenty-one 
days. This was not so with the Dutch rules in any single season, unless dates 
before July were excluded. If we consider July in relation to the tempera- 
ture-humidity rule, we find that blight followed within fifteen days in four 
seasons out of ten and within twenty-one days in six seasons out of ten 
(Table i^). This is very similar to the results from the Dutch rules. 
Examination of column 8 shows, however, that fairly good forecasts could 
have been made every year except in 1932, 1934 and 1937 when there 
would have been no forecasts. 

A comparison of the general results of the application of the two sets of 
rules indicates that the Dutch rules give rather too many critical periods 
and the temperature-humidity rules too few. Might not a slight modifica- 
tion of the temperature-humidity rules lead to their wider application? It 
is possible, for example, that a period of one or two hours of slightly lower 
relative humidity than 75% would not check the effect of the two-day 
period in bringing about the production of conidia. When the weather 
data available for 1931--9 are re-examined to allow for this, the total 
number of critical periods is increased from nineteen to twenty-eight, and 
accurate forecasts can be made of the slight outbreaks in 1934 and 1937 not 
previously forecasted. The percentage of critical periods witliin the fifteen- 
day period, out of the total observed before the outbreak, is, however, only 
slightly greater in this wider application, i.e. fourteen out of twenty-eight 
(50%) instead of eight out of nineteen (42 %), and for the twenty-one-day 
period sixteen out of twenty-eight (57 %) instead of eleven out of nineteen 
(58 %). The advantage of a slightly wider interpretation of the humidity 
rule is therefore negligible. 

The important exception of the June outbreak of 1931 may now be 
considered. Humidity and temperature were both favourable on 28, 30 
and 31 May, and on 5, 6, 8, 9, 10 and ii June, i.e. on nine days out of 
fifteen. In no other year were there more than four such days in this 
period. Again, the number of days in May 1931 with the requisite tem- 
perature and humidity was six: in other years the number ranged from 
none to three. If we examine the Dutch rules for the same period there 
is no such outstanding indication: the number of favourable days before 
10 June, the date of outbreak, was only three. There were also three days 
before this date in 1933 and two in 1929, 1932 and 1935. Also in May 1931 
and in six other seasons there was only one favourable day according to the 
Dutch rules. 

It may thus be concluded that from 1929 to 1939 both the Dutch rules 
and the temperature-humidity rule gave useful indications of blight in 
South Devon, and that the temperature-humidity rule was slightly more 
precise and simpler to apply. 

Application of the temperature-humidity rule to other districts 

W^st CornwalL Conditions in West Cornwall are particularly interesting 
because the potato crop consists of early varieties put in the ground very 
early and often dug before blight has appeared in other parts of England. 

Weathet and Outbreak of Potato Blight, A, Beaumont 51 

Limited crop rotation and close planting favour the progress of the disease. 
It might therefore be expected that the conditions preceding a blight out- 
break might correspond less closely to the rules than in South Devon. 
Investigation has shown that this is not so, but that, as will be seen in 
Table 2, the rules apply to West Cornwall equally satisfactorily. 










Table 2. Potato blight in West Cornwall 1931-9 

Date of 

Number of critical 
days preceding 
outbreak of blight 


Up to 15 days Over 15 days 


12 May 



30 May 



7 May 



16 May 



Very local 







20 May 






Crop lifted early 

o o 

The data in the table for 1931 and 1932 were prepared from records 
taken at Gulval, which is in the centre of the early potato area. Early 
afternoon readings of humidity were not taken after 1932 and the data for 
the remaining years were obtained from the Daily Weather Map, which 
gives the minimum temperature at Penzance and the 13 hr. relative 
humidity at the Lizard. These are less satisfactory, as the minimum 
temperature is often slightly lower at Penzance than at Gulval, and at the 
Lizard there is generally a higher humidity. To correct this a minimum of 
80 % humidity was taken for the table. 

The records show that blight followed the first critical periods within 
fifteen days in three seasons out of the four in which blight occurred in 
the district. In the fourth season (1933) blight followed thirty-five days 
after the first critical period and two days after the second. There was no 
critical period and no blight in 1936 and 1939 ; and in 1934, 1935 and 1938 
the first critical period came too late, most of the crop having been lifted 
before blight made its appearance. 

In 1931 the critical period coincided with the outbreak of blight, which 
was probably governed by the weather of 28 and 29 April and 5, 6 and 7 
May, when the slightly lower minimum temperatures that prevailed on 
those dates, coupled with the high humidity, were still sufficient to bring 
about blight. This was also noticed for earlier seasons by Wiltshire (1931). 

France, The temperature-humidity rule was tested in France in 1938 by 
Limasset (1939). Observations at Versailles showed that there was no 
favourable period until September, the only days following the rules being 
26 July, 23 August, and 20, 25, 26 and 27 September. Spread of blight 
began early in August but was slight; and as was found for Newton Abbot 
in 1934 and 1937, the method was not successful in forecasting the start of 
a slight epidemic. As pointed out by Limasset, under such conditions the 
microclimate determines the extent of the disease in any given locality and 


52 Transactions British Mycological Society 

cannot be represented adequately by a sii^igle set of observations. At the 
end of September, when the epidemic became more extensive, the rules 
could be correctly applied. 

Other observers in the Hautes Pyrenees in 1938 found that the outbreak 
of blight (20 August) followed a favourable period from 15-18 August in 
which the rules were obeyed (Ministere d’ Agriculture, 1939, p. 114). 


A detailed study has been made of the meteorological data recorded at 
Seale-Hayne Agricultural College, Devon, during the eleven years 1929- 
39, in relation to outbreaks of potato blight, and it has been shown that the 
Dutch rules governing the date of the blight outbreak also apply in South- 
west England. Their practical value as a basis for forecasts is lessened by 
the fact that favourable days occur which are not soon followed by blight, 
so that the application of the rules in Devon in every season except 1931 
indicated blight earlier than it appeared. During the period in question 
blight appeared in June only in 1 93 1 . Excluding this year and considering 
only the July and August weather data it was found that blight followed 
the first critical date within fifteen days in three seasons out of ten and 
within twenty-two days in six seasons out of ten. The rules provided no 
explanation of the early outbreak of 1931. 

It is difficult to obtain the necessary data for the application of the Dutch 
rules, and a simpler modification has been devised and called the * tem- 
perature-humidity rule’. The conditions are (i) minimum temperature not 
less than 50° F. and (2) relative humidity over 75 % for two consecutive 
days. This rule has been found to give good results in Devon and West 
Cornwall most seasons. Blight followed the first critical dates within 
fifteen days in three seasons out of eleven and within twenty-two days in 
five seasons out of eleven. The data also throw light on the exceptionally 
early outbreak of 1931. Excluding this year and considering only the July 
and August weather records it was found that blight followed the first 
critical period within fifteen days in four seasons out of ten and within 
twenty-two days in seven seasons out of ten. In the remaining three 
seasons there was no critical period. This failure to indicate blight was not 
a disadvantage from the practical point of view because the blight 
epidemic was so slight in those seasons. 

On the whole the temperature-humidity rule is regarded as slightly 
more useful for blight forecasting than the Dutch rules. 

Weather and Outbreak of Potato Blight. A. Beaumont 53 


BeaumonTj a. (i929~32)* Dep. PL Path. Seale^Hayne agric. ColL 

Beaumont, A. (1930), Rep, Dep, PL Path. Seale-Hayne agric, ColL p. 22. 

Beaumont, A. (1940). Potato blight and the weather. Trans, Brit. mycoL Soc, xxiv, 266. 

Crosier, W. (1934). Studies in the biology of Phytophihora infestans (Mont.) de Bary. 
Mem, Cornell agric. Exp. Sta* no. 155, 

Dufri^noy, J. (1933). In Rapports sommaires sur les travaux accomplis dans les labora- 
toires en 1932. Ann, ^piphyt, xix, 1-46. Abstract in Rev. appL Myc, xiii, 76. 

Limasset, P. (1939). Recherches sur le Phytophihora infestans (Mont.) de Bary. Ann. 
jSpiphyt, n.s. v, 2i~39. 

Melhus, I. E. (1915), Germination and infection with the fungus of the late blight of 
potato. Bull, Wisconsin agric. Exp. Sta. no. 37. 

Meteorological Office (1928). The Book of Normals of Meteorological Elements of the 
British Isles. Section VI. Normals of Relative Humidity. 

Ministj^re d’Agriculture (1939). Comptes rendus sommaires des Travaux des Stations et 
Laboratoires de Recherches Agronomiques en 1938. 

Naoumova, N, a. (1935). [On forecasting the appearance of Phytophihora infestans on the 
potato.] PL Prot. Leningrad, nr, 51 -4. Abstract in Rev. appl. Myc. xv, 522. 

Van Everdingen, E. (1926), Het verband tusschen de weergesteldheid en de aardappel- 
ziekte [Phytophihora infestans). Tijdschr. PlZiekt. xxxii, 129. Abstract in Rev. appl. Myc. 
V, 627. ■% 

Van Everdingen, E. (1935). Het verband tusschen de weergesteldheid en de aardappei- 
ziekte (tweede mededeeling). Tijdschr. PlZiokt. xli, 125-33. Abstract in Rev. appl. 
Myc. XIV, 715. 

Van Poeteren, N. (1928). Een waarschuwingsdienst voor het optreden van de aardap- 
pelziekte. Versl. PlZiekt. Dienst Wageningen, no. 53. Abstract in Rev. appl. Myc. xi, 95. 

Van Poeteren, N. (1931). Verslag over de werkzaamheden van den Hantenziekten- 
kundigen Dienst in het jaar 1930. Versl. PlZiekt. Dienst Wageningen, no. 64. Abstract 
in Rev. appl. Myc. ya, 

Wiltshire, S. P. (1931). The correlation of weather conditions with outbreaks of potato 
blight. Quart. J. R. met. Soc. lvii, 304-16. 

[Accepted for publication 30 October 1946) 

[ 54 ] 


By JOHN GRAINGER, Department of Plant Pathology^ 
West of Scotland Agricultural College^ Auchincruive^ Ayr 

(With Piate V and 7 Text-figures) 

Corn mildew, caused by the fungus Erysiphe graminis DC., can assume 
serious economic proportions in the west of Scotland, when cereal crops are 
grown for green fodder. It causes some damage on grain crops, but 
probably develops too late in the growing period to curtail the production 
of grain very seriously. 

Studies on the ecology and biology of E*. graminis in various countries 
have given somewhat discordant results, and it is probable that different 
strains are involved on the same crop in different regions. There are, of 
course, strains upon various grasses and cereals which will not inter-infect. 
The strain chiefly used in the present experiments appears to attack only 
oats; it will not infect wheat. These investigations were planned to acquire 
information as to the factors responsible for the severity of attack by E. 
graminis in the west of Scotland. They evaluate some aspects of the major 
factors of climate, soil nutrition and composition of the host plant. 
Literature on E. graminis has been adequately reviewed by Gherewick 

The effect of climate 

Preliminary correlations of climate and the severity of corn mildew^ 
suggest that the fungus owes more to increase in relative humidity (r.h.) 
than to increase in summer temperature, in the severity of its attack 
(Table i). Records showed, moreover, that the r.h. associated with 
the heaviest infection of mildew (greenhouse, Table i), reached 100% 
on six days out of the forty-two covered by the preliminary inquiry. The 
R.H. of the other two sites never attained saturation at 9 a.m., when the 
readings were taken. 

Table i. General climatic relations o/* Erysiphe graminis on oats 


Plant Pathology fields, Auchincruive 
Oat trials, Auchincruive 
Greenhouse, Plant Pathology Dep. 

Mildew % of 
leaf area 




During development of disease 

. . A ■ 

i ' \ 

Av. temp, ® F. Av. % r.h. 

59 79 

69 77 

59 88 

It is suspected that the disease becomes severe when atmospheric 
humidity reaches saturation for appreciable periods. That this idea has 
some foundation in fact is shown by Text-fig. i, which integrates the 
number of hours of ioo % r.h. per week with the percentage of leaf area 

Ecology of 'Etysi^ht gTdm 55 

covered by the fungus. The humidity was computed from a continuous 
hygrograph record; the infected leaf area was estimated by comparison 
with ^standard area’ diagrams (see Appendix). The year 1944, with a large 
number of hours of 100% r.h. per week allowed a development of the 
fungus to cover 50 % of the leaf area; 1945, with fewer hours of saturation 
per week, had a much lower infection of about 5 %. These figures refer to 
the general level of mildew in the oat crops sown round the meteorological 
station at the Department of Plant Pathology, Auchincruive. 

Text-fig, X. Relation between the intensity of Erysiphe graminis and the number of hours lOO % 
R.H. per week. Vertical columns represent the number of hours per week with saturated 
atmosphere (left-hand vertical scale). The full black line shows the development of mildew 
(right-hand vertical scale). 

Measurements of spore germination in atmospheres maintained at 
standard relative humidities confirm this dependence of the fungus upon 
atmospheric saturation. Conidia from mildewed oat leaves were shaken on 
to sterile cover-slips, which were stored in sealed glass tubes above solutions 
containing varying concentrations of calcium chloride. A drop of water 
containing conidia was placed on a cover-slip in each series, and this was 
similarly stored over water to prevent evaporation^ The whole series was 

56 Transactions British My cological Society 

placed iix an incubator at 22° C. Several repetitions were made, and the 
collected results are given in Table 2. Spores of the fungus Cladosporium 
herb arum are always mixed with those of Erysiphe graminis in natural 
infections, and their differing responses to r.h. are noteworthy. 

Table 2. Germination 0/ Erysiphe graminis conidia 

Storage conditions 

Maximum percentage germination 
Erysiphe graminis 

Germination, Cladosporium herbarum 
Solution to give storage conditions 

% f »H. 














+ + 

+ + 















Spores of the oat strain of E. graminis under test germinate much 
better in air at 100% r.h. (PL Va) than in actual water, whilst 
germination does not take place at relative humidities of 85 % or less. 
Spores of Cladosporium herbarum germinate copiously in water and 100% 
R.H., and grow well at relative humidities down to 76 %, but not at 66 %. 
Spores of Erysiphe graminis did not grow beyond the stage shown in PI. Va 
in these germination tests, but it is noteworthy that Cladosporium herbarum 
grew and produced chains of conidia in water, and over all relative 
humidities down to 76 %. 

It is reported from Canada (Cherewick, 1944) that conidia of the oat 
strain of Erysiphe graminis will germinate at 100 % r.h. and also at o % r.h. 
over barium perchlorate. The Scottish strain clearly has not this wide 

E. graminis will attack young oat plants twenty days after germination, if 
the atmosphere is saturated for sufficient periods. It will also grow upon 
old leaves, as long as they are green, but ceases to produce conidia when the 
foliage turns brown. This is apparently not related to the degeneration 
and loss of the chlorophyll, for no significant germination was obtained 
when conidia of E, graminis were dusted over drops of oat-leaf extract, of 
green sycamore-leaf extract and anthocyanin-containing red sycamore- 
leaf extract. Oat-leaf extract, when filtered through filter-paper, gives 
a cloudy suspension of chlorophyll; sycamore leaves provide a clear, 
opalescent green or red fluid when similarly prepared. Chlorophyll and 
anthocyanin therefore appear to be without effect on the germination of 
Erysiphe conidia. It is also interesting that Oxyacanthae (DC.) de 

Bary was found growing on white leaves of hawthorn at Auchincruive in 
1945 ; 

Mildew is often found more plentifully towards the base of a cereal crop, 
than among the upper foliage. This presumably reflects the higher r.h. 
nearer the ground. It is, indeed, frequently found that humidity is highest 
near the ground, (e.g. in wheat, Text-fig. 2), or in the lower parts of the 
crop (e.g. oats in Text-fig. 3). These layers presumably reach atmospheric 
saturation before the air above, and retain it for a longer period. An 
interesting comparison was possible between two crops of oats grown near 
one another at Auchincruiye in 1946. One crop (P in Text-fig. 3) was 

Height above ground in ft 

Height above ground in ft. 

58 Transactions British Mycological Society 

planted near the top of a low hill, and had an average of i % mildew. The 
other was 300 yards to the south-east, and lay in a slight hollow, to leeward 
of the hill in relation to the prevaihng wind. It experienced much lower 
wind velocity (A, Text-fig. 3), with about 3 % more r.h. than the other 
crop, and also had 25 % mildew. 

It would seem, from general observations, that corn mildew is generally 
more severe in the west of Scotland than in most parts of England. Heavy 
attacks occur in aU regions, but when I moved from Yorkshire to the west 
of Scotland I obtained a strong impression that mildew is much more 
generally severe in the latter region. Data compiled from the Air Ministry’s 
Monthly Weather Reports (1938-44) for the various meteorological districts of 
the British Isles show that the west of Scotland has a large number of rain 
days per annum, and a slightly higher average r.h. than most regions of 
England. Differences of average r.h. are not large, but are fairly significant 
(e.g. ^ = between 0-05 and 0-02 by #-test when comparing the west of 
Scotland (83%) with the midlands of England (78%)). The west of 
Scotland, moreover, has significantly higher average monthly hunudities 
from May to August, when cereal crops make most growth, than districts in 
England. Figures of r.h. during the summer months for the west of Scotland 
are only exceeded by the north of Scotland, where mildew is also severe. 

These field and regional studies all tend to confirm the relation between 
severity of Erysiphe graminis and a high r.h. They also point to the need for 
a more detailed and comparative estimation of disease intensity between 

The effects of major soil nutrients 

Most workers report increasing severity of mildew when nitrogenous 
manures are added to the soil. Field experiments at Auchincruive in 1945 
and 1946 suggest that phosphate reduces the severity of mildew and potash 
increases it. These experiments were carried out in conjunction with 
Mr John McFarlan of the Department of Agriculture, West of Scotland 
Agricultural College. The results are shown in Tables 3 and 4. 

They show that both nitrogen and potash tend to make mildew more 
severe, while phosphate reduces its severity. The effect of nitrogen has been 
confirmed by adding varying amounts of nitrogen to water cultures of oats 

Additions of phosphate to land which is already well supplied with 
phosphate appear to be without effect upon the severity of mildew, when 
oats are grown (Table 6). It is probably only the correction of actual 
phosphate deficiency which will reduce the .severity of mildew. 

Various practical indications suggest that continued applications of 
artificial manures have an adverse effect upon the severity of mildew. 
This can be seen by comparing the plot totals for ‘no addition’ and 
‘complete manuring’ in Tables 3 and 4. The question has also been tested 
by solution culture, using two methods : 

(1) Varying the concentration of solution directly (curve A in Text-figs. 
4, 5 and 6). 

(2) As (1), but the solution renewed every fourteen days (curve B). 


Ecology of ’Exjsvght gxsjokas. J. Gr airier 

Table 3. Soil factors and Erysiphe graminis, 1945 

(Phosphate-deficient soil. Peel Hill, Auchincruive) 
Mildew, % of 

Treatment leaf area Plot totals 

No addition 






Nitrogen 52 

No nitrogen 




Phosphate 17*5 

No phosphate 


Nitrogen, phosphate 


Potash 41 

No potash 


Phosphate, potash 


No addition 4 

Complete manuring 


Nitrogen, potash 


Nitrogen, phosphate, 



Table 4. Soil factors and Erysiphe graminis on Victory Oats^ 1946 

(Soil low in phosphate, Donald’s Thorn, Auchincruive) 

Mildew % of 
leaf area 

Treatment Block I Block 11 Plot totals 

No addition 








1 5 Nitrogen 226 

No nitrogen 171 



25 Phosphate 180 

No phosphate 222 

Nitrogen, phosphate 


1 8 Potash 2 1 0 

No potash 182 

Phosphate, potash 


17. No addition 37 

Complete manuring 55 

Nitrogen, potash 



Nitrogen, phosphate, 




Manures applied at rates per acre, in 1945 and 1946 


3 cwt. 

Nitro chalk 


6 cwt. 

18 % superphosphate 


2 cwt. 

60% muriate of potash 

Table 5- 

. JVitrogen 

level and Erysiphe graminis on Star Oats 

Mildew % of leaf area 

Culture solution 

average of all leaves 

Sachs’s + micro-solution 

: with ^ normal N 


with normal N 


with 2 times normal N 


with 4 times normal N 


I Table 6* Phosphate level and Erysiphe graminis on Star Oats 

Original cont. 

mg./ioo g. 

Mildew % of 

of dry soil 

leaf area 

Soil A 


No addition 


-h Superphosphate 




No addition 


4- Superphosphate 


Soil G 


No addition 

, . : 5 

(phosphate fixing) 

4- Superphosphate 


Each pair of soils was compared separately, and at different times, so that the amounts of 
mildew recorded are only comparable as between no addition and the application of super- 
phosphate in each case, 

1 V : V , ■ . ' 

Tillei's per plant Dry wt. per plant in g. Mildew, % of leaf area 

Text-fig. 4. Disease incidence of Erysiphe graminis at different levels of inorganic nutrition in 
solution culture. -4= original solution retained throughout. ^5= solution changed every 
fourteen days. JV on the horizontal axis is normal Sachs’s solution with micro addition. 

Text-fig. 5. Data for dry weight and tilleririg of Star oats grown at different levels of inorganic 
nutrition in solution culture. 4 —original solution retained throughout, solution 
changed every fourteen days. # on the horizontal axis is normal Sachs’s solution with 
micro addition. 

Text-fig. 6. Data for total carbohydrate, nitrogen and ash at different levels of inorganic 
nutrition in solution culture. .4= original solution retained throughout. j5= solution 
changed every fourteen days. JSf on the horizontal axis is normal Sachs’s solution with micro 
addition. Percentages of nutrients are expressed on a dry-y^eight basis. 


Ecology of gmmmis. J. Grai^^ 

Sachs’s solution with additions of micro-elements was made up in seven 
concentrations, from one-eighth normal to eight times norm'aL Five culture 
jars were set up for each strength. Results of dry weight and of tillering 
are given in Text-fig. 5 and of the incidence of mildew in Text-fig. 4. They 
show that changing the solution every fourteen days [B) resulted in in- 
creased dry weight and increased tillering, as compared with the retention 
of the same solution throughout (T). Mildew was much more severe in 
series .5 (Text-fig. 4), where the solution was changed every fourteen days. 
The incidence of mildew also increased with increasing concentration in 
series .4 (Text-fig. 4) up to twice the normal concentration. This provides 
contributory evidence that raising the level of inorganic manuring brings 
increased incidence of mildew. 

The effect of composition of the host plant 

Differences between the amounts of mildew with varying treatment in the 
last experiment seemed to provide the possibility of a good test as to 
whether the amounts of carbohydrate, Kjeldahl nitrogen and ash present 
; in the host had any effect upon the severity of mildew. 

5 All the five plants of each treatment were accordingly dried at 98° C. 

and were subsequently analysed for total carbohydrate by the Willaman 
I and Davidson (1924) method, for nitrogen by Kjeldahl technique, and for 

f ash by incinerating in a muffle furnace. The results are portrayed in 

' Text-%. 6, which shows that no relation appears to exist between the 

^ amounts of carbohydrate and ash and the level of disease (compare with 

Text-fig. 4). Total nitrogen is low in series which has severe mildew, and 
: is higher in series with much less disease. This is rather surprising, in 

ii view of the increase in severity of Erysiphe with unbalanced nitrogenous. 

I manuring. It suggests that the increase of disease by nitrogenous manuring 

is brought about rather by an indirect effect of the nitrogen upon the 
anatomy of the plant, than by its amount within the host. 

Another method of testing consists ip growing plants under varying lengths 
of day. It was noticed that spring-sown wheat being grown under eight- 
hour day conditions developed mildew, whereas the same crop under 
normal illumination did not do so, nor did autumn-sown wheat under any 
condition of illumination in 1945. This material was also examined for 
total carbohydrate, nitrogen and ash, with results set forth in Table 7. The 
strain of mildew was not that used in the foregoing experiments, but the 
results are nevertheless useful. They show very little difference in carbo- 
hydrate content between the four treatments, but there is more ash and also 
more nitrogen in the plants which developed mildew. The association of 
higher nitrogen with severe disease is in contrast with the results of the last 
experiment, and it was thought that the higher nitrogen might reflect the 
delay in ripening of the spring-sown, short-day plants. That phenology is 
not the whole explanation of attack by mildew appeared in 1946, when the 
disease again developed on short-day oats and wheat, but not so severely on 
the plants grown with normal length of day, except that infection of the 

62 Transactions British Mycological Society 

wheat became more general in August. This appearance of the disease 
occurred in the early spring, when the autumn-sown wheat had leaves 
only a few inches long. Mildew is also recorded on wheat in June and on 
wheat and oats in August. The oats had green leaves in both short-day and 
long-day settings in August, so there is here no question of ripeness pre- 
venting attack by mildew. The collective results shown by Text-figs. 4 and 
6 and Tables 7 and 8 do not furnish any correlation between the severity of 
attack by Erysiphe graminis and the major constituents of the host. 

Table 7 

Total % 

% total 





Date Height Total d.w. 



of fig. ft. in. 



in leaves % ash 


LD AS Sq. Master 

2 1 June 3 2 






White Victor 

22 June 3 3 







24 June 3 I 






LD SS Sq. Master 

29 June 2 loj 






White Victor 

30 June 2 9 







4 July 2 8 






SD AS Sq. Master 

r8July 2 3 






White Victor 

20 July 2 3 







20 July III 






SD SS Sq. Master 

* 10 






White Victor 

* 11 







* 7 




10*37 . 


Samples for analysis were taken 20 August 1945 

* Did not flower 

Table 8 

% mildew 

% total 


% , — 


carbo- % 


Date of flg ft. in. 

solids 10/3 


5/8 hydrate ash 


Star oats LD SS 

i6July 2 3 

23*5 — 



41*6 10*0 



— II 

21*7 — 



42*0 8*2 


Atle wheat LD AS 

24 June 2 0 

27*1 0 



39-8 7*9 



— 16 

24*9 10 



42*0 4*6 


WHte Victor LD AS 

24 June 2 0 

29*1 0 



38*9 7*9 


wheat LD SS 

15 Aug. I 10 

22*3 — 



40*9 7*8 



— I 9 

25-3 15 



36*2 11*9 


— SD SS 

— 6 




28*6 12*0 

■■ 3*85 

Samples for analysis were taken 5 August i 


LD Long day (normal variation in length of day) 

SD Short day (eight hours exposure to light) 

AS Autumn sown (Table 7) 33 Sept. 1944, (Table 8) 14 Nov. 1945 
SS Spring sown (Table 7) 28 Feb. 1945, (Table 8) 12 Apr. 1946 
— Not sown on 10 March 1946 

It is also interesting that mildew on oak, wild cherry and rose does not 
appear to be correlated with the amounts of total carbohydrate, or with the 
content of reducing sugars (Table 9). These results are from material taken 
for analysis on 1 1 August 1936. 

Ecology of Erysiphe graminis. J. Grainger 


Table 9 


Solids carbohydrate 
% of fresh wt. % of dry wt. 

A. Microsphaera Alni on oak {Quercus robur). Dean Nick, Huddersfield 
Healthy young leaves 29*25 38*93 

Mildewed young leaves 34*57 14*71 

Healthy old leaves 41*70 i 3*45 



% of dry wt. 




B. Podosphaera Oxyacanthae on wild cherry {Prunus padus). Dean Nick, Huddersfield 

Healthy young leaves 28*74 19*85 

Mildewed young leaves 28*29 19*52 

Healthy old leaves 32*28 10*00 

G. Sphaerotheca pannesa on cultivated rose, Huddersfield 

Healthy young red leaves 24*5 34*2 1 

Healthy young green leaves 36*4 25*78 

Green-red leaves with mildew 30*0 26*62 

Green-red leaves; mildew removed 28*7 20*12 

Healthy old leaves, i 37*9 16*88 

Healthy old leaves, 2 26*5 30*95 











A strain of Erysiphe graminis which attacks oats in the west of Scotland has 
been mainly used in the present investigation. Conidia germinate much 
better in a saturated atmosphere than they do in water. Germination is 
low at 90 % R.H. and apparently does not take place at lower values of r.h. 
Spores of Cladosporium herbarum which accompany those of Erysiphe graminis 
will grow and fruit in water and at humidities down to 76 %, but not 

Climatic correlations suggest that the severity of the disease in the field is 
related to the degree of saturation of the atmosphere. Regional and field 
studies support this view. Severity of the disease appears to owe much 
more to the inorganic nutrition than to the gross composition of the host. 
Practical control of the disease in the west of Scotland lies in the avoidance 

of excess nitrogen and potassium in the soil, the correction of any deficiency 
of phosphate, and the avoidance of heavy and frequent application of 
artificial manures. 

The writer wishes to express his grateful thanks to members of his staff 
at the Department of Plant Pathology, Auchincruive, for their general 
assistance, and to Mr H. F. Dovaston for the photomicrographs of Plate IV. 
Mr John McFarlan has maintained valued collaboration with the field 
experiments, and the determinations of Table 9 were made in 1937 with 
the help of Mr A. Broadbent of Huddersfield. 


In this paper the severity of mildew has been expressed as a percentage of 
the total leaf area of the host which was covered by the fungus. This has 
been accomplished conveniently by comparing all leaves of the plant with 
‘standard area^ diagrams (Text-fig. 7). An average ofallthese readings gave 

The standard area’ (hagrams used for the determination of percentage leaf area 
hy ETysiphe graminis. 

Ecology of Btysiphc gmrm^ 65 

the disease measurement for the plant as a whole. In field measurements it 
was found adequate to assess the disease intensity at the base, middle and 
top of the crop, the average of all three being the reading for the sampling 
station. Four stations were estimated on plots acre in area, nine in 
acre plots, whilst twenty sampling stations were measured in each field, 
when it was necessary to make more widespread estimations. 

The ^standard area’ diagrams were made by spacing ovoid dots upon 
a leaf-shaped area, the dots having standard percentages of the leaf area. 
They thus provide a standard of comparison for different persons, for 
different regions and for different times. It does not seem to n.atter very 
much that the standard areas can never equal those of any leaf in actual 
shape and distribution; the human eye detects fairly readily the similarity 
of distribution frequency and pattern. 


Cherewigk, W. J. (1944). Studies on the biology of Erysiphe graminis DC. Canad. J. Res, 
XXII, 52-86. 

WiLLAMAN, J. J. & Davidson, F. R. (1924). Some modifications of the picric acid method 
for sugars. J. agric. Res. xxviii, 474-88. 


A (above) . Germination of conidia of Erysiphe graminis in air at 100% r.h. B (below). Conidia 
in air at 90 % r.h. Only one spore has germinated, and many spores are shrivelled. 

[Accepted for publication 2 November 1946) 

[ 66 ] 



Department of Cryptogamic Botany^ The University, Manchester 
(With Plates VI and VII and i Text-figure) 

Since certain strains of Penicillium notatum had been reported to be very 
variable and much variation had been observed in the cultures used for the 
commercial production of penicillin, it was decided to investigate this 
phenomenon. The strain examined most thoroughly was N.R.RX. 1249 
B2I3 which was distributed by the Northern Regional Research Labor- 
atories, Peoria, Illinois, and has been widely used for commercial production 
by the surface culture method. It was derived by successive selections 
from the original Fleming strain. Raper and Alexander (1945) have 
described this strain and given an account of its history. They reported the 
occurrence of paler, less highly sporulating variants and of darker, more 
heavily sporulating variants. The present investigations were begun before 
the publication of Raper and Alexander’s paper, and where the observa- 
tions coincide the results obtained are in agreement with those of Raper 
and Alexander. Variants showing differences in sporulating capacity 
have been obtained by single conidial platings and by streaking from 
mycelial patches on old cultures. The poorer sporulators were found to be 
the higher penicillin yielders up to a certain limit. Very poor sporulajtors, 
however, seem to be very poor yielders of penicillin as reported by Raper 
and Alexander (1945). 

It is the purpose of this paper to deal in greater detail with the occurrence 
of variations affecting sporing capacity, with a view to elucidating the 
cause or causes of the variation. 


Streaking from old, cultures 

Variants were obtained from patches of mycelial growth overgrowing the 
normal conidial type on old cultures. By streaking and finally inoculating 
on to plates of plain agar it was usually possible to obtain single hyphal 
isolations of these types. These gave apparently pure cultures of mycelial 
habit. Cultures obtained in this way I have called 'mycelial types’. 
Similar 'mycelial types’ were also obtained by conidial platings of old 

cultures. , ... , . , . 

oingte conidial isolations 

Variants were also obtained by plating out suitable dilutions of conidial 
suspensions taken from cultures of various ages. The conidial suspensions 
were made in physiological saline and detergent (o-i % cafeolene oil). The 
suspensions were well shaken to break up chains of conidia and filtered 
through absorbent cotton under sterile conditions. This removes pieces of 
mycelium and clumps of conidia. The density of a conidial suspension was 

m Penicillium notatu^ 67 

found by making haemocy tome ter counts and from this suitable dilutions 
were estimated and plated. The haemocytometer observations also per- 
mitted a check on the presence or not of clumps of conidia. If such were 
observed the suspension was shaken and re-filtered. This was very seldom 
necessary, the conidia being usually well separated. A small amount 
(o-2 c.c.) of the required spore dilution was spread over the surface of an 
agar plate with a spreader made of bent glass tubing. It was preferred to 
spread the spores over the surface of the agar rather than to mix them in 
cooled melted agar and then pour them, since the former method gave 
a more regular germination of spores which facilitated a comparison of the 
single spore colonies. 

With some exceptions the medium used was 3 % malt extract with 2 % 
agar. On this medium the types of variants studied were clearly differ- 
entiated from each other and the parental type. 


An ampoule of Penicillium notatum N.R.R.L. 1249 B21 (circulated by the 
Northern Regional Research Laboratories, Peoria, Illinois), was opened 
and subcultured. The subculture was transferred to plain agar and single 
hyphal isolations made. These proved to be of two types, indicated here as 
T I and T2. Type i (probably the parental 1249 B21 type) was paler and 
less heavily sporulating than T2. It is not known whether the colour 
difference between the two types is entirely due to difference in conidia 
production or not. T2 also differed from T i in that it produced less 
yellowing of the medium. 

On malt agar T i gave a smooth velvety growth, pea green (PL 47, 
Ridgway colour chart) on the eighth day. T 2 gave a similar type of growth, 
sage green (PL 47) on the eighth day. On the modified Czapek agar used by 
Raper and Alexander, when three cofcnies were inoculated on one plate, 
the edge of the plate was reached by the tenth day. T2 showed a rather 
fiat velvety growth with a few shallow radial furrows. T i was similar to T 2 
with slightly more pronounced furrows. The colours on the tenth day 
according to the Ridgway colour chart were artemisia green (PL 47) at 
the outside, shading to Hathi grey (PL 52) in the centre for Ti, and 
artemisia to celandine green at the edge, shading to dark olive grey in the 
centre for T2. In both.cases the colony reverse was yellow. 

Colonies of each type were more uniform and more distinct from the 
other type on malt than on Czapek agar. 

At first it was not known whether T i or T 2 was the standard 1249 ^ 

type. However, according to the description given by Raper and Alexander, 
Ti would appear to correspond to the original type, T2 probably being 
a variant derived from it. 

Conidial platings were made from the original single hyphal isolates, and 
single conidium cultures were obtained by transferring a piece of agar con- 
taining a single germinated conidium on to an agar slant under the dissecting 
microscope. Spores from the single spore cultures were in time plated out 
and in this way data were obtained concerning the number of variant 
colonies given by single hyphal and single conidium cultures. 


68 Transactions British Mycological Society 

Table i . Production of pale and dark variants by H. T. {single hyphal) 
and S.S. {single conidium) cultures of Ti type 
















: 1 1 days 




















28 „ 






— . 




42 » 










10 „ 










10 „ 










18 „ 





















13 days 









4 » 









21 „ 










10 „ 






























Total H.T. and 
S.S. cultures 










Table 2. Production of pale and dark variants by H.T. {single hyphal) 
and S.S. {single conidium) cultures of Ti type 


Total % % Other other Total % 











: 10 days 










10 „ 










10 „ 





— ■ 






II s, 





— - 





18 „ 









■ — 

28 „ 











II » 










42 „ 



— . 








42 „ 









; 1*5 












4 days 








10 „ 




— . 


- — 



13 « 






— . 















21 „ 









I* I 


29 M 











29 » 








* — 

" ' — ' 











Total H.T. and 
S.S, cultures 









* Obtained from cultures used in 

production at 

the I.C.I. plant 

in Trafford Park. 

The results obtained from cultures of T i type are shown in Table i and 
those from darker types in Table 2 . These tables include a number of dark 
and light selections obtained from cultures used in production at the I.C.I. 
plant in Trafford Park. The cultures within one group were not identical in 
appearance and it is possible that they differed by certain genetic modifying 
factors. Table I shows the results obtained by plating seven single hyphal 
and five single conidium cultures of Type i. There were no statistically 
significant differences between platings from the single hyphal and single 

Spontaneous Variation in Penicillium notatum 69 

conidium cultures. There were 0*8% of darker variants, 1*5 % of paler 
variants and o*i % of other variant types, 2*4 % in all. 

The darker types (Table 2) gave only o*6 % of variants, 0-2 % of darker 
variants and 0-4% of paler variants. The cultures of T i gave approxi- 
mately four times as many variants as those to T2. This spore plating 
method for detecting variants might be expected to favour the higher 
sporulating types but since this would tend to increase the number of 
darker variants and decrease the number of paler variants recorded in 
both T I and T2, it is possible that the darker types are actually more 
stable than the paler ones. It is also possible that the cultures included in 
Table 2 are not homogeneous with respect to rate of variation, some being 
more stable than others. Very stable strains have been obtained from T i. 
On one occasion a very dark heavily sporulating type giving excessive 
yellowing of the medium was obtained from a culture of T i . This proved to 
be very stable and paler variants derived from it were also relatively stable. 

The preliminary investigations showed a remarkable degree of uni- 
formity both in the proportions of mutants and in their appearance. It 
seemed possible therefore that the greater part of the variability observed in 
this strain might be due to the constant occurrence of a particular type of 
change rather than to random variations of different origin. In order to 
test this hypothesis T i was studied more intensively. 

Analysis of Type i variants 

From a plating of T i, a standard T i, a pale variant T i a^ and a dark 
variant colony T i were streaked and finally single hyphal isolations were 
made. Conidia from these isolates were plated and single conidium 
isolations were made. It was found that T i <2, gave a still paler variant 
type, T I also variants resembling T i and T i b. This seems to 

indicate that changes in sporulating capacity occur in steps since Tia gives 
a still paler variant Tia^ and that these changes are reversible since Tia 
gives a type closely resembling T i. The results from seven single conidium 
cultures of T i are given in Table 3, and from eleven single spore cultures of 
Ti^, in Table 4. It will be noticed that every culture gave colonies of 
a paler type, Ti^ and that all except two gave a darker type, Ti b. The 
percentages of these variants were remarkably constant from one culture to 
another, with the exception of S.S. 3 which was recorded as having a visible 
pale sector and which gave a higher percentage of pale variants than its 
sister cultures. The average percentage of T 1 6 variants was 0*4 and of 
T i a, excluding the culture with a visible pale sector, 2*2. The percentage of 
variants of all types was 2*7, excluding S.S. 3. Single conidium cultures of 
the pale type T i ^ gave 0-9 % of still paler variants, T i a^, 1*7 % of variants 
resembling T I and 0*3 % of variants of T i type, as shown in Table 4. 
Again the percentages of variants of the different types were very constant 
among the offspring of the sister cultures. Only in one culture, which had 
been recorded as having a visible green sector, was there any striking 
deviation from the average percentages. The average percentage of total 
variants, excluding the culture with a sector, was 3* o, not significantly 
different from the 2 '7 observed in the case of T i . Only four cultures of 

70 Transactions British Mycological Society 

Table 3. Production of dark {Tib) and pale {Tia) variants by 
sister single conidium cultures of Ti type 











colonies variants T i ^ 








S.S, : (1)12 days 


— ■ 






(2) „ „ 









* (3) » .. 






: — 




(4) SJ 33 










(5) 33 33 



— . 






■ '1*4 ■■ : 

(6) 19 33 










(7) ,3 ,3 










(8) „ ,3 















■ 3*5 





Total excluding 
S.S. (3) 











Recorded as having a 

visible pale sector. 

Table 4. Production of dark {Ti and Tib) and pale ( Ti fig) 
variants by sister single spore cultures of Ti a type 

Total T I 
colonies variants 



Tib % 
variants T i ^ 

T I % 

variants T itZ2 





S.S.: (I) 

12 days 

















— ■ 






















T -3 



































1 1 




















* ( 9 ) 














































Excluding S.S. (9) 



1 18 1*7 22 0*3 65 0-9 

Recorded as having a visible dark sector. 



Table 5. Production of darker ( Ti b^ variants by cultures of Tib type 








H.T. (42 days) 



. — 

S.S. (i) (14 days) 




S.S. (2) (14 days) 




S.S. (3) (14 days) 






Table 6. Production of darker {Tia, Ti and Tib) 
variants by cultures of Tia^ type 


T I a 




Ti^ % 




colonies variants Tia variants 


variants Ti^ 



S.S. (i) 40 days 






2 ’ 3*9 


^ 5*7 

S.S. (2) II days 






7 3‘i 









9 3*2 



Tib were tested (Table 5). Two of them gave no variants and the other 
two gave variants of a still darker type, T i h^. The average percentage of 
these mutants was o-8 % but all except one came from one culture so this 

Spontaneous Variation in'? tmc^Xmmn^ 71 

percentage may be higher than that which would be obtained if more 
cultures were tested. The darker variant T i has not been further in^ 

Two cultures of type Ti^2 have been plated and each gave variant 
colonies o^T la^ T i and Tib types. T i <^2 appears to be the most unstable 
type so far encountered since it gave an average of 18-9% of variant 
colonies. However, since T la^ produces very few conidia, variants with 
greater sporulating capacity would be expected to produce more conidia in 
proportion to the amount of mycelium present. Hence the conidial 
plating method of testing variation might be expected to exaggerate the 

Text-fig. I. Variation of Ti and derivatives. 

actual variation. However, sectoring was more frequent for Tia^ than for 
the other types, therefore it is probably actually less stable than the other 
members of the series, although probably not to the extent indicated by the 
percentage of variant colonies obtained. 

The production of variants by the different types is illustrated in 
Text-fig. I. 

In discussing the tables, it has been assumed that the variants occurring 
in different cultures were similar variants. 

An attempt was made to see how far these variants of similar appearance, 
occurring regularly in different single spore cultures were identical. 
Streaks were made from variant colonies, Conidial suspensions were made 
from these streaks and inoculations were made at three places on an agar 
plate as follows, using the paper disc method of inoculation devised by 
Dr P. H. Gregory. Two small sterile filter paper discs were immersed in the 
spore suspension and placed side by side on the agar plate. Two discs from 
a suspension made from a similar type of different origin were placed at 
a second position and one disc from each suspension was placed at the third 
position. Variants of Ti^» type arising from four different Ti a type 
cultures were tested against a variant of Ti ^ type arising directly from T i. 
In some cases two different colonies from the same type were tested. In 

72 Transactions British Mycological Society 

addition, two T 1 6 cultures arising in two different single conidium cultures 
of T I type were tested against the control. 

In all these tests the three colonies on one plate were identical in 
appearance (see PL VI, fig. 3) . , 

Three variants of T i type from three different single spore cultures of 
T I ^2 type were similarly tested and gave identical colonies on each plate. 
See (PL VI, fig. 4). 

Variants of Ti^ type obtained from four different single conidium 
cultures of T i type were combined with a T i type from a fifth single 
conidium culture. Mass inoculations were used instead of spore suspensions 
because of the reduced sporulation in the type. The five cultures tested 
appeared identical (see PL VI, fig. 5). 

Variants of Tia^ from four different single conidium cultures of T i 
type tested in the same way appeared to be identical (PL VI, fig. 6). 

This method, of inoculating cultures of similar appearance side by side, 
was used, as being the most stringent test of identity available, in the 
absence of sexual reproduction. Besides affording the possibility of com- 
paring the cultures, in conditions as nearly as possible identical, it also 
permits of heterokaryort formation at the line of junction of the two types. 
The use of heterokaryons is at present the only possible test for allelism in 
Fungi imperfecti. When two different mutants form a heterokaryon it often 
happens that the mutation in each locus is recessive to its allele on the 
other nucleus, so that the heterokaryon is different in appearance from its 
two components. If, for example, two T i types of different origin 
inoculated side by side gave a green streak at their line of junction it could 
be deduced that the two types, while phenotypically similar, were geno- 
typically different. However, the absence of such an effect does not prove 
that they are identical although it is in accordance with such a view. 

Description of T 1 variants 

Macroscopic, A description is given of the variants on malt agar which was 
used as the principal medium and also on the modification Czapek agar 
used by Raper and Alexander (1945). The various types in the Ti series 
are most clearly differentiated on malt agar on which they give a smooth 
velvety growth. T i and T i b show a completely fiat growth but Tia shows 
very shallow radial furrowing, and T i zzg a more distinct radial furrowing. 

follows : 


8 days on malt 


Sage green 29'"', 

PL 47 


Pea green 29"", 

PL 47 


Greenish glaucous 
33 "^ H. 41 


Sea-foam yellow 25", 

II days on malt 
Pea green 29"'', PI. 47, 
shading to celandine 
green PL 47 

Court grey 29"", 

PL 47 

Greenish glaucous 
33^'^ to yellowish 
glaucous 25''', PL 41 
Sea-foam yellow 25'L 
PL 31 ; 

10 days on Gzapek-Dox 
Celandine 33"" through 
artemisia 33 PL 47, 
to deep olive 23''''' or 
mouse grey PL 


Artemisia 33^"'j PL 47, 
at edge shading to 
Hathi grey 35'''^''' or 
storm grey 35^'"', PL 
52, in centre 

Spontaneous Variation in Penicillium notatum 73 

It will be noticed that colour differences can be observed among the 
different types on malt agar. On Czapek agar, however, the colours are 
the same for T i, Ti a, T i ^2? but the central paler regions differ in size, 
being largest in T i <22 and smallest in T i. The diflFerences in macroscopic 
appearance seem to be due to the fact that Tib ->T i ->T i a i form 
a series of types tending to spore later and less freely. 

Microscopic, Preparations were made by fixing portions of mycelium in 
formol-alcohol for a few minutes, staining in chlorazol black E (saturated 
solution in 70 % alcohol), rinsing in 70 % and mounting in gum chloral. 
Chlorazol |)lack was used as a fungal stain by Armitage (1944). The use of 
gum chloral as a mountant was suggested by Dr P. H. Gregory. It has an 
advantage over balsam and gum damar for this material in that de- 
hydration and clearing in xylol is eliminated. Hence a greater number of 
penicilli are left, comparatively undisturbed. It is not known how long 
such preparations will last but they have been kept for six weeks without 
appreciable fading. The material was taken from nine-day cultures except 
in the case of T i ^2 • (I^ 'this type no penicilli could be found in nine-day 
cultures so mounts were made from 26-day old cultures.) 

T I. Raper and Alexander (1945) have given a detailed description of 
the conidial apparatus of 1249 B21 and it is therefore unnecessary to 
repeat this here. So far as I can determine, T i corresponds to 1249 
and the description given for this strain holds for T i. Metulae vary from 
one to four in number, sterigmata from one to five with three as the most 
frequent number and a certain number of irregular penicilli are present 
(P 1 .X,%. 8 ). 

Tib (PI. VII, figs. 7, 1 1) forms more conidia than T i and this is correlated 
with differences in the number of sterigmata and in the number of conidia 
to be found on one sterigma. Thus the most frequent arrangement of 
metulae was in groups of four as compared with three in T i . Some 
sterigmata were observed with chains of eight conidia attached whereas the 
greatest number of conidia attached to one sterigma in T i was six. It is 
realized that the actual number of conidia observed in the chains is likely 
to be affected by the mounting process but it is felt that the observations 
give a fair representation of the relative lengths of chains in the different 

In Ti^i (PI. VII, fig. 9) the greatest number of sterigmata 
observed had only one conidium and the longest chains recorded consisted 
of only four conidia. Most frequently only one sterigma was present, but 
perfectly regular penicilli with groups of four sterigmata were observed. 
However, irregular penicilli were much more frequent in this type than in 
the two preceding types; Ti^ had 1/30 irregular penicilli, Ti, 5/42 and 
Tifli 14/36; 

The irregularities consisted chiefly in the formation of secondary metulae 
and in a spreading in the position of the sterigmata. The lower-sporulating 
types tend to have much less compact penicilli than the normal and the 
various abnormalities seem to be related to this tendency. 

Ti % (PL VII, figs, 10, 12) shows a still further reduction in sporulation 
capacity than T i a. More penicilli consisting of only one or two sterigmata 


Transactions British Mycological Society 

were present than in Ti a, A characteristic of this variant is that the 
sterigmata are often flattened. The variation in spore size is much greater 
than in the other types. 

Mycelial types. As previously described^, patches of mycelial growth 
frequently occur on old cultures of P, notatum. Such patches have been 
observed on cultures started from a single conidium and therefore origin- 
ating from one nucleus. A number of mycelial types have been oi^tained 
from single conidium cultures by streaking and finally making hyphal tip 
inoculations. Several such cultures have been isolated from single spore 
cultures of similar type. These cultures differ from each other in general 
appearance, in the amount of pigment produced and in colony shape but 
they are all alike in that they rarely revert to the conidial form. The change 
from the conidial to the mycelial habit appears to be essentially irreversible. 
Most mycelial types produce some conidia and it has twice been possible 
to observe that these conidia are formed on greatly reduced, almost 
abortive penicilli. However, when these conidia are plated out they produce 
colonies of mycelial type only. In other characteristics, such as colony 
shape, the mycelial types are subject to variation. All the mycelial types 
tested by the surface culture method have been found to be poor yielders of 
penicillin: this is in accordance with the results of Raper and Alexander. 
They did not distinguish between mycelial types which could not revert to 
conidial types and very poor sporulating types which could throw higher 
sporulating types but it is probable that their ‘g’ and 'M’ types are 
mycelial types in the sense of this paper. Mycelial types have been 
obtained from both Ti and T2 strains. It is probable that mycelial 
patches may differ in frequency and extent in different strains but few 
quantitative data have been obtained on this point. 

Mycelial types can also be obtained by single conidium isolations from 
old cultures. One particular mycelial type with a spiky transparent type of 
growth has been obtained in this way from four different single spore 
isolations of T i and from one of T 2. One mycelial type, obtained on an 
old T I culture, after continuing unchanged through a number of mass 
transfers eventually gave a sector of this peculiar transparent type. 


The evidence presented indicates that T i and its derivatives are subject to 
frequent variation of a particular type. The variation is progressive since 
Tia gives a paler type Ti^g, and T16 gives a darker type; and it is 
reversible to a certain extent since Tia gives the parent type Ti. The 
frequency with which T i b types are recovered from T i is 0*4 % and from 
T I 0-3 % indicating that the change to T i b occurs at the same rate in 
T I and T i a. The reversion from T i a to T i probably takes place at the 
same rate as the original change (T i to T i a) , since T i gives 2-2 % of T i 
types and T i a gives 1 7 % of T i type. T i ^2 appears to be more variable 
than Ti and Tia-^ giving 18*9 % of variant colonies. sectors 

more frequently than the other^^^^t^^ it is probably, actually, as well as 
apparently, lesk stable than T i and T i a. 

Spontaneous Variation in Penicillium notatum 75 

The question arises as to whether the changes are to be considered as 
being mutations or not. In so far as they are spontaneously occurring 
changes which are transmitted from cell to cell, they fall within the 
category of mutational changes. Since sexual reproduction has not been 
observed in this fungus it is not possible to follow the transmission of these 
changes during sexual reproduction. We cannot determine in this way 
whether the changes are transmitted by the nucleus. It is difficult however 
to see how results of such regularity could be obtained if the, factors 
causing them are cytoplasmic. It may be possible to get some evidence 
regarding the location of these factors by studying diploid strains. ‘ Gigas’ 
forms of a different strain of N.R.R.L. 1978B have been obtained by 
camphor treatment. These forms are probably diploid. It is hoped to 
obtain such ‘ gigas ’ forms of 1249 B21 and study variation in them. If the 
forms are diploid and if the changes which affect sporulating capacity 
occur in the nucleus^ the variation pattern would be different from that of 
the haploid. 

. Mutation rate 

It is clear that if the variants observed are due to mutations at a particular 
region (region being used in a general sense to mean either a gene or 
a piece of chromosome) the changes are of such a high frequency that the 
region may be considered a 'mutable region’. Such a mutability may be 
due to faulty reproduction of a particular segment leading to repeated 
reduplication and loss. If this is so, conditions affecting the rate of nuclear 
division might possibly affect the mutation rate. It is hoped to test the 
effect of environmental conditions such as changes of temperature on the 
rate of mutation in an attempt to obtain some clue as to the fundamental 
nature of the mutation. 

The determination of the mutation rate is a very difficult problem. 
Delbriick (1945) has discussed the subject in relation to bacteria and much 
of his discussion is relevant to fungi also. So far as the radial growth of 
a colony on an agar surface is concerned it is clear that at any one point 
on the circumference of the colony there is very great competition as to 
which hyphae shall continue the growth and produce a new sector. When 
mutant nuclei are present in fewer numbers than the original nuclei, they 
have less chance of being present in the hyphae which carry on the growth 
of the colony. Mutants will only succeed in establishing themselves as 
sectors when they occur with a very high frequency or when they have an 
appreciably higher growth rate than the original type. So far as the aerial 
growth is concerned, the conditions have not been clearly worked out. 
However, it would be seen that if there is any weighting it would be in 
favour of the type present in greatest numbers, i.e. the original type. 
Therefore ’we may consider the percentage of mutant types recovered as 
a lower limit for the actual mutation rate. The actual mutation rate is 
probably not lower than the percentage of mutations recovered and is 
possibly higher. 

Bunting (1940^, b) describes the occurrence of colour variation in Serratia 
marcescens which is similar in certain respects to the variation described in 

76 Transactions British Mycological Society 

this paper. She found progressive variation in the sense that the dark red 
type gave a bright pink variant which in its turn gave a pale pink variant. 
She also found reversion since the bright pink and pale pink types both 
gave dark red types. This corresponds to the fact that T la and T i <22 both 
give T I types. However, in the case of Serratia the dark red types gave 2 % 
of bright pink colonies whereas the bright pink colonies gave 42 % of dark 
red colonies. 

Skovsted (1943) has described what he calls 'successive mutation’ in 
the yeast-like species, Nadsonia RichterL Mutants were picked out as sectors 
reaching the edge in giant colonies. A number of single-celled cultures of 
the same original type each gave the same two types of mutants. Each of 
these types produced three new types. Two of these types were tested and each 
gave four new types. Five of these eight types were tested and gave altogether 
twenty-three new types. Thus although only two new types were obtained 
from the original t^e, many new types were derived from the mutants. 
Skovsted did not observe any examples of reverse mutation. However, the 
competition for survival in a colonial form is such that sectors can only be 
formed by mutants which occur very frequently and have a growth rate not 
less than the normal or by mutants which have a higher growth rate on the 
normal. The shape of the sectors illustrated by Skovsted shows that many of 
the mutants have a growth rate greater than that of the normal. In such 
conditions reverse mutations would not be normally detected even if they 
occurred because they could not compete with the new type. It might be 
possible, however, to detect them by altering the medium so as to favour the 
original type at the expense of the mutant. They could also be detected, if 
frequent, by plating out a large number of single cell colonies. Until such 
experiments have been made one cannot be sure that reverse mutations do 
not occur in Madsonia, 

Mycelial types 

Mycelial types have been described for Penicillium notatum by Hansen and 
Snyder (1944). They relate the occurrence of such types to the 'dual 
phenomenon’ which they have described in a large number of fungi 
imperfecti. Single spore (multinucleate) isolations of a number of fungi 
gave three types: M, mycelial, C, conidial and MC, intermediate. M types 
gave only M types, C types gave only C types, whereas MC types gave all 
three types when single spore isolations were made. AfC types could be 
synthesized by putting M and C types together. MC types were thus found 
to be heterokaryons containing Af and C nuclei. Later it was found that M. 
types could arise from C types and that this often happens with great 
regularity whereas M types are stable, never having been observed to 
revert to C types although subjected to different cultural conditions. 
Earlier, Mohendra and Mitra (1930) described the occurrence of segrega- 
tion of black colony (conidial) and white colony (mycelial) types during 
spore formation in Sphaeropsis malorum. This appears to be an example of the 
' dual phenomenon ’ of Hansen ( 1 938) and of Hansen and Snyder ( 1 948) and 
here also the mycelial type did not revert to the conidial type, Mohendra 
and Mitra relate the supplantation of the black colony type by the white 

Spontaneous Variation in Vcmcilliv^ 77 

colony type during mass conidial transfers to the slower germination and 
reduced viability of the black colony types. 

Mycelial types have been observed mostly in fungi imperfecti and 
therefore it has not been possible to test the segregation of such types during 
sexual reproduction. However, Hansen and Snyder (1943) were able to 
cross M and C strains of Hypomyces Solani f. Curcurbitae. It was found that the 
mycelial-conidial character segregated in a i : i ratio and was independent 
of the incompatibility factors. It was also found that whereas the conidial 
type was hermaphrodite the mycelial type did not produce perithecia and 
could only function as a male. Hansen and Snyder therefore consider the 
change from the conidial to the mycelial habit to be a true mutation from 
a hermaphrodite to a male condition. In Bombardia lunata^ Zickler (1937) 
found the lanata factor, which appears to be a factor for mycelial growth, to 
be absolutely linked to a female ^sex realisator’ and to be independent of 
the incompatibility factors. Robbins and Ma (1945) describe physiological 
experiments on Trichophyton mentagrophytes in which they used the conidial 
wild type and a series of ‘pleomorphic’ types obtained from old cultures of 
the conidial type. The situation in Trichophyton resembles that in Penicillium 
in that a number of such types are found and in the non-reversion of these 
types to the conidial habit. 

The mode of origin of these mycelial types is an interesting problem. The 
orthodox genetical hypothesis would be that they were mutations to a type 
or types which could grow on the surface of old cultures and which would 
tend to be selected in transferring from old cultures. However, such 
mycelial types would not be expected to become established in the wild 
because of the absence of a satisfactory propagating mechanism such as 
that provided by the conidia on the wild type. In Hypomyces Solani f. 
Cucurbitae and Bombardia lunata^ where the presence of a sexual stage permits 
the testing of the mutation theory, the mycelial type behaved as a single 
gene change, but in Penicillium and Trichophyton no such direct test is yet 
possible. If the mycelial types are mutational in origin, the question arises 
as to whether the different mycelial types within one species are due to 
different mutations or whether there is one mutation for mycelial types and 
the differences are due to modifying factors or cytoplasmic effects. The 
presence of the mutable series affecting sporulation capacity suggests that 
mycelial types may occur as an end result of this series. A frequently 
occurring type of mutation such as this would account for the frequent 
appearance of mycelial patches on old cultures. If this is so and all mycelial 
types are due to one type of change then such types when inoculated 
together would not give the wild type. This was reported by Pontecorvo and 
Gemmell ( 1944 ) in Penicillium and has been confirmed by me ( 1 946 ) . Also T r a 
and T I when grown together do not give patches of T i type, which is in 
accordance with the view that they are in some sort of allelic series. These 
results however maybe due to the less highly sporulating habit being dominant 
over the conidial habit as suggested by Pontecorvo and Gemmell ( 1944 ). 

It is hoped that further investigations will disclose whether there is a 
connexion between the ‘mutable region’ and the production of mycelial 
types or not. 


Transactions British Mycological Society 

Penicillin production 

In these experiments as in those of Raper and Alexander (1945) there 
was a negative correlation between sporulation capacity and penicillin 
production in the conidial types. This, however, may be due to some 
physiological or mechanical effect of sporulation on penicillin production 
rather than to a direct effect of the gene affecting sporulation on penicillin 
production. Mycelial types were found to be poor yielders. 

A series of variants with increased production of yellow pigment showed 
a decreased production of penicillin. This accords with previous reports. 


The standard strain of 1249 B21 constantly throws paler and darker 
variants. The paler variants throw the darker variants, revert to the 
parental type and throw still paler variants. The latter are unstable giving 
all the darker types. 

The colour is related to, and may be conditioned * by, sporulating 

The greater part of the variation exhibited by young cultures is due to 
changes in this particular "mutable region h 

Patches of mycelial growth frequently occur on old single spore cultures 
as well as on old mass inoculation cultures. Mycelial types which do not 
normally revert to the conidial type can be isolated from these patches. 

It is suggested that there may be a relationship between the "mutable 
region’ and the production of mycelial types but no direct link between the 
two phenomena has yet been observed. 

It gives the author great pleasure to thank Imperial Chemical Industries, 
Ltd. for financial support. Prof. C. W. Wardlaw for constant help and 
encouragement and Mr E. Ashby for taking the photographs. 


Armitage, F. D. (1944), Chlorazol black E as a stain for mycological specimens. Trans. 
Brit. myc. Soc.'KKvii, 131-3. 

Bunting, M. I. (1940 a). A quantitative study of the dark red to bright pink variation in 
Serratia marcescens. J 5 ar^. xxxnc, 15. 

Bunting, M. I. (1940^). A description of color variants produced by iS'^rra^fa marcescens. 
J. Bad. xxxiXj 108, 

Deubruck, M. (1945). Spontaneous mutations in bacteria. Ann. Mo. Bot Odn. xxxi^ 

Hansen, H. N. (1938). The dual phenomenon in imperfect fungi. Mycologia^ xxx, 442— 
55 - 

Hansen, H. N. & Snyder, W. C. (1943), The dual phenomenon and sex in Hypomyces 
solanii. cucurbitae. Amer Rol xxx, 419— 22. 

Hansen, H. N. & Snyder, W. C. (1944). Relation of dual phenomenon in Penicillium 
to pehicillin production. Science, N.S. xcix, 2570, 264-5. 

Mohendra, K. R. & Mitra, M. (1930). On the cultural behaviour of Sphaeropsis 
malorum. Aa. Rat, Xa«fif., XLrv, 175, 541-55, 

Spontaneous variations in Penicillium notatum 79 

PoNTEGORVOj G. & Gemmell, A. R. (1944). Genetic proof of heterokaryosis in Penicillium 
notatum. Nature^ Lond., ciisr^ 

Raper, K. B. & Alexander, D, F. (1945). Penicillin. V. Mycological aspects of 
penicillin production. J. Elisha Mitchell Sci. Soc. lxi, 74-113. 

Robbins, Wm. J. & Ma (Roberta) (1945). Growth factors for Trichophyton mentagro- 
phytes. Amer, J. Bot. xxxn, 509-23. 

Sansome, E. R. ( 1 946) . Induction of ‘ gigas ’ forms of Penicillium notatum by treatment with 
camphor vapour. Nature^ Lond.^ CLvn, 843. 

Skqvsted, A. (1943). Successive mutations in Nadsonia Richteri Kostka. C.R. Lab. 
Carlsberg^ Serie physiol, xxiii, 409-53. 

ZiCKLER, H. (1937). Die vererbung des Geslechts bei dem Ascomyzeten Bombardia 
lunataZckl. indukt. Abstamm.- u. VererbLehre, 'L' kx.vui, 40^-8. 

Plate VI 

All cultures photographed on malt agar 

Fig. I. Seven-day colonies from a plating of Ti^z type. Three variants of Ti type and two of 
T I ^ type are visible. 

Fig. 2. Seven-day colonies from a plating of T i type. One variant of T i ^ type and eight of T i 
type are visible (dark colonies on right due to shadow) . 

Fig. 3. Inoculation of one Ti ^ strain at top of Petri dish, inoculation of a second Ti b strain of 
independent origin at left of Petri dish and side-by-side inoculations of two strains at the 
right of the Petri dish. One strain shows a mutant sector. 

Fig, 4. Similar inoculations of two strains of T i type separately and combined (combination at 
the right) . 

Fig. 5. Similar inoculations of two strains of Tia type. 

Fig. 6. Similar inoculations of two strains of T i ^3 type. 

Figs. 3-6. Eight-day cultures. 

Plate VII 

Photomicrographs, figs. 7-10 x 500; figs, n, 12 x 1000 

Fig. 7. T I ^ from nine-day culture. 

Fig. 8. T I from nine-day culture. 

Fig. 9. Tia from nine-day culture. 

Fig. 10. T I from twenty-six-day culture. 

Fig. 1 1 . Single penicillus of T i 

Fig. 12. Single penicillus of T 1^2* 

Slides fixed formol alcohol, stained chlorazol black and mounted gum chloral. 

{Accepted for publication 2 November 1946) 

[ 80 ] 



The Freshwater Biological Laboratory ^ Wray Castle^ Ambleside^ and 
Department of Botany^ Birkbeck College^ London 

(With Plate VIII and 3 Text-figures) 

Since Myzocytium megastomum was originally described by De Wildeman 
(1893) a number of references to it have appeared in the literature 
(Scherffel, 1914; Skvortzow, 1925; Cejp, 1935; Berdan, 1938; Karling, 
1942), but because of the lack of knowledge of the zoospores, its exact 
systematic position has remained in doubt. According to Sparrow (1943), 
this species has not been recorded from Britain. However, Prof. C. T. 
Ingold (personal communication) in 1944 found this organism attacking 
Euastrum ansatum^ Closterium costatum^ C. rostratum and Pleurotaenium Ehren- 
bergii from Woodhouse Eaves near Leicester (Text-fig. 2 a-f)^hut again the 
zoospores were not observed. In July 1946 I found Myzocytium megastomum 
^diVd.dtizmg Spirotaenia condensata Breb., in Little Green Tarn, Claife 
Heights, near Hawkeshead, Lancashire, and the biflagellate zoospores 
were observed, showing that this fungus really belongs to the genus 
Myzocytium. A single specimen of what is believed to be this fungus was 
found in Closterium from the Clay Pond, Wray Castle, Windermere, in 
September 1946. It differs from the material from Green Tarn and 
Woodhouse Eaves in the more globular zoosporangia (2 1-26 yu in diameter) 
and in the larger zoospores (9 /x* in diameter) . Mature sporangia and stages 
in the formation of the biflagellate zoospores are shown in Text-fig. 3, and 
PL VIII, figs. 3, 4. Young thalli and resting spores were not observed. An 
account of Myzocytium megastomum on Spirotaenia condensata follows. 

The young endobiotic thallus is relatively short and not subdivided by 
constrictions. At this early stage of development the empty encysted 
zoospore and its germ tube apparently connected with the thallus, could 
usually still be seen outside the host cell (Text-fig. i b) . The thallus elongates 
and becomes constricted into portions each of which later becomes a single 
sporangium. The young sporangia are often separated from one another by 
a plug of refractive material (Text-fig. i a) which disappears as the 
sporangial wall thickens. Such plugs were not observed by Prof. Ingold in 
the material from Woodhouse Eaves (Text-fig. 2 b, c). The number of 
sporangia from any one infection into which the thallus is transformed 
varies from about one to eight, but more than one thallus may be found in 
a single host cell. The mature sporangia are ovoid, ellipsoidal or spherical 
(32-19*8 ju, long X 15-13*2 /x diam.)^^ hyaline cytoplasm containing 
many scattered highly refractive globules (Text-fig, i {;) . 

By theTime the sporangia are mature the host chloroplast is reduced to 
a brown residue, although it still exhibits a somewhat spiral form. At last, 
from each sporangium a discharge tube is formed which is expanded 

Text-fig, I. Myzpcytium megastomum, chain of young sporangia separated by refractive plugs, 
X 500. very young thallus ; zoospore case, and infection tube visible outside the host cell, 
X 975. eleven mature sporangia, the host chloroplast now very shrivelled; germinated 
zoospores of a chytridiaceous organism also present, x 750. early stage in the develop- 
ment of the discharge tube from the sporangium, X975. discharged undifferentiated 
zoospore mass, x 750. /, zoospore mass showing differentiation of zoospores and fringe of 
flagella, x 750. g, a zoospore, X975. empty sporangia with typical swelling of the 
discharge tube within the host cell, x 750. 7, a cell containing two dehisced 

sporangia, and three resting spores; the males are distinguished by the absence of a discharge 
tube, X 50a Cosmarium contractum with possible dwarf thalli, x 500. the same in 
Staurastrummmtumix^oo, . 

82 Transactions British Mycological Society 

immediately within the host wall. This is the characteristic feature of this 
species. According to the place of germination of the sporangia relative to 
the host cell wall, the endobiotic swelling may either be almost sessile on 
the sporangium (Text-fig. i d, h) or at some distance from it (Text-fig. 2 d,f). 
The discharge tube, except for the intramatrical swelling (4-6 p, in 
diameter), is equally cylindrical throughout its length (2-6 p wide) and 
may extend up to 30 p in length outside the host wall (Text-fig. ih). The 

Text-fig. 2. Myzocytium megastomum, a, empty zoospore case with germ tube on Closterium 
costatum; chain of sporangia each with a conspicuous vacuole in Closterium rostratum; c, chain 
of sporangia in Pleurofaenium Ehrenbergii; d, germinating sporangia, each discharge tube 
swollen immediately inside the algal wall, in Closterium costatum ; e, four empty sporangia in 
Euastrum ansatum; f, chain of sporan^a in various stages of development in Closterium 
costatum. In most of the figures the disintegrating host contents are omitted. (Drawn by 
G.T. Ingold.) 

extramatrical prolongation of the discharge tube ih the material from 
Woodhouse Eaves appears to be relatively short (Text-fig. 2e,f). On 
deliquescence of the apex of the discharge tube, the contents of the 
sporangium emerge to form a spherical granular mass lyp in diameter 
with numerous small refractive globules (Text-fig. i e). This mass undergoes 
slight amoeboid movements and, some ten minutes later, the zoospores 

Myzocytium megastomum. Hilda M. Canter 


gradually become differentiated, and the flagella appear as a fringe of 
short, actively waving structures around its periphery (Text-fig. i/) . The 
zoospores remain entangled by their flagella for some time, but finally 
break free and swim away individually. 

Each zoospore mass becomes resolved 
into ten to fourteen somewhat bean- 
shaped zoospores (4'5-“5 long x 5*6- 
7^ diam.) with granular protoplasm 
containing several small refractive 
globules. There are two flagella of 
about equal length inserted laterally 
in a slight depression. One flagellum 
is directed backwards and the other 
forwards when swimming (Text-fig. 


Resting spores are formed abun- 
dantly together with the sporangia, 
and agree with those described by De 
Wildeman ( 1 893) . The mature resting . 
spores are spherical, 13-15/x in di- 
ameter, with a smooth thick wall, and 
oily contents (Text-fig. ij). No con- 
vincing young stages in the develop- 

ment of these resting spores were seen, T V ’rST"/ 

but it is clear that they are formed by 
a sexual process. In only one specimen 
was any definite fertilization tube con- 
necting the male and female game- 
tangia observed (Text-fig. ij). The 
male gametangia can always be dis- 
tinguished from the empty sporangia 
by the lack of discharge tubes. The 
resting spores are usually formed at 
the side of the female gametangium 
nearest the male. Their germination 
was not observed. 

Young thalli of a chytridiaceous 
organism were also present on the 
Spirotaenia cells (Text-fig. ir), but 
further stages in the life history of this 
fungus are not yet known. 

The following records of 
megastomum hsNQ already been made: 
from Belgium, Switzerland and 
Norway (De Wildeman, 1893, 1895, 

1896); Hungary (Scherffel, 1914); 

Manchima. (Skvortow, 19=5)^ Bo- 

hernia (Gejp, 1935); and America ctomm sp. X300. 


84 Transactions British Mycological Society 

(Berdan, 1938). Karling (1942) rightly lists Ancylistes miurii Skvortzow 
(1925, p. 432, figs. 7“io) as a synonym. 

From the many observations on Myzocytium megastomum it would seem 
that the expansion of the discharge tube within the host wall is a good 
specific distinction separating M, megastomum from the closely allied M. 
prolifemm (Schenk) . If this is so, then, the M. proliferum of Martin ( 1 92 7 ; in 
Cladophora sp.), the specimen recorded by Sparrow (1943; in Closterium 
costatum^ Farlow Herbarium no. 642), and that of De Wildeman (1895, 
p. 76, pi. 2, figs. 7-9; in Euastrum) should be included in Myzocytium 
megastomum^ a course not followed l3y Karling (1942). De Wildeman’s 
specimen in Euastrum is peculiar in producing only one sporangium instead 
of a chain of sporangia, and probably represents a reduced form. Similar 
simplified thalli were seen in the Leicestershire collection (Text-fig. 2 e); 
by myself in August 1946 from the plankton of Lake Windermere, South 
Basin, in Cosmarium contr actum (Text-fig. ik) and Staurastrum lunatum 
(Text-fig. im), and by Petersen under Myzocytium irregulare (1909; 402, 
fig. i 6 d; 1910; 538). Karling (1942), in agreement with Fischer (1892), 
De Wildeman (1896) and Minden (i9iiy5 has suggested that Bicrium 
transversum and Bicrium naso (Sorokin, 1883) may also represent dwarf 
thalli of Myzocytium^ and since B, naso has an endobiotic swelling on the 
discharge tube this species would be referable to M. megastomum- The 
superficial similarity of these simplified forms of M. megastomum^ in the 
smaller desmids, with Olpidium immersum Sorokin cannot be overlooked, 
and since the zoospores of 0. immersum have not been observed this may be 
found to belong to the genus Myzocytium- However, only when the zoo- 
spores, and resting spores of dwarf thalli of Myzocytium, Bicrium naso and 
Olpidium immersum have been observed, and inoculation experiments on 
various desmids been carried out, will the true nature of these fungi be 

My thanks are due to the Director of The Freshwater Biological Associa- 
tion, Wray Castle, Windermere, for the use of a laboratory in which this 
work was done, and especially to Prof. G. T. Ingold for helpful criticism, 
and permission to publish his figures of Myzocytium megastomum- 


Berdan, H. (1938). Revision of the genus xxx, 396--415. 

Gbjp, K. (1935). The parasites of Conjugates in Bohemia, Illrd contribution. Bull. 
Int. Acad. Sci. BoMme, xlv, i~i2. (Separate.) 

Fischer, A. (1892). Phycomycetes. Die Pilze Deutschlands, Oesterreichs und der 
Schweiz. Rahenhorst. Kryptogamen-FLi{/^yi—/^'^o. Leipzig. 

Karling, J. (1942). The simple holocarpic biflagellate Phycomycetes. New York City. 

Martin, G. W. (1927). Two unusual water moulds belonging to the family Lageni- 
diaceae. Mycologia, 'Km, 

Minden, M. von ( i 9 i i ) . Chytridiineae, Ancylistineae, Monoblepharidineae, Sapro- 
legniineae. Kryptogarmnfl. Mark Branderiburgy v, pt. 3, pp. 353-496. 

Petersen,. H. E. (1909). Studier over Ferskvands-Phycomyceten. Bidrag til Kund» 
skaben om de submerse Phycomyceters Biologi og Systematik, samt om deres 
Udbredelse i Danmark. Bot Tidsskr. xxix (4)? 345-440. 

Myzocytium megastomxim. Hilda M. Canter 85 

Petersen, H. E. (1910). An account of Danish freshwater Phycomycetes, with bio- 
logical and systematical remarks. Ann. MycoL^ BerL^ viii, 494-560. 

ScHERFFEL, A. (1914). Kisebb Kozlemenyek a kryptogamok korebol (Kryptogamic 
Miszellen). Bot. KozL 

Skvortzow, B. W. (1925). Zur Kenntnis der Phycomyceten aus der Normandshurei, 
China. Arch, Protistenk, li, 428-33. 

Sorokin, N. W. (1883). Aper^u systematique des Ghytridiacees recoltees en Russie et 
dans FAsie Centrale. Arch. Bot Nord, Fr, ii, 1-42. (Issued as a separate.) 

Sparrow, F. K. (1943). Aquatic Phycomycetes. Ajin Arbor, U.S.A. University of Michigan 

WiEDEMAN, De (1893). Notes mycologiques. II. Ann, Soc, Beige Micro, (Mem.) xvn, 

WiLDEMAN, E. De (1895). Notes mycologiques. IV. Ann, Soc, Beige Micro, {Mem,) xix, 

WiLDEMAN, E. De (1896). Notes mycologiques. VII. Ann, Soc, Beige Micro, {Mem,) xx, 

Myzocytium megastomum De Wildeman 

Fig. I . Part of a Spirotaenia cell containing sporangia. The disorganized spiral chloroplast of the 
host is clearly visible, x 780. 

Fig. 2. Three empty zoosporangia and a resting spore in Spirotaenia : {a) the male, {b) the female, 
containing a thick-walled resting spore, x 1 230. 

Figs. 3, 4. Parts of a Closterium cell from the Clay Pond, Wray Castle, with zoosporangia in 
various stages of development. The swelling of the discharge tube immediately within the 
host wall is well marked. X450. 

{Accepted for publication 5 Movember 1946 ) 

[ 86 ] 


By W. C. M.OO'K'Ej^ Plant Pathology Laboratory^ Harpenden 

1. Peronospora Dipsaci TuL, in C.R. Acad. ScL, Paris, xxxviii, 1103 (1854); 
Gaumanu, Monogr. Gattung Peronospora, 238 (1923). 

On. leaves of Dipsacus sylvestris L., Dundry, Somerset^ May 1941 (L. 
Ogilvie) and Bricketwood, Herts, October 1946 (A. Smith). 

2. Sphaerotheca fiiliginea (Schlecht.) Salm., in Mem. Torrey hot. CL xxix 

On cultivated Doronicum, Wye, Kent, July 1938 (W. M. Ware); Luton, 
Bedfordshire, October 1939. Probably widely distributed. 

In the material from Luton the perithecia were 81 “96 jjl (average 87 ft) in 
diameter, the asci 62-72 x 5i--55ft (average 67 x 52 ft) with 6-8 ascospores. 

3. Oidium sp. 

On Antirrhinum mqjus L., Bucks, March 1928 (W. Buddin); Devon, 
Newton Abbot, April 1928 (A. Beaumont); Cambridge, February 1934 
(F. T. Brooks) ; and later at Cardiff (J. Rees) ; Shrewsbury (N. C. Preston) 
and Harpenden. 

Usually slight, under glass, and probably not uncommon. At Harpenden 
seen also after planting out, and at Shrewsbury severe on bedded out plants. 
At Harpenden forming a sparse white web or dusty covering on under- 
surfaces of leaves, and occasionally on upper surface. Gonidia hyaline, 
almost rectangular, with slightly rounded ends or occasionally barrel- 
shaped, 24-45 X 1 2-1 7 ft (average of forty spores 30 x 14-5 ft). 

4. Oidium sp. 

On soft young tips of gorse shoots {Ulex sp.) in the Malvern Hills, July 
1931 (G. H. Pethybridge). 

The only previous record of a powdery mildew on Ulex appears to be one 
in Ireland in 1906. Adams (1906) t tcov 6 .t& Cicinnobolus Ulicis n.sp. as 
‘parasitic on one of the Erysiphaceae on stems and leaves of Ulex europaeus, 
which it covers with a felt of hyphae bearing pycnidia (Figs. 2, 3 and 4), 
July. On Great Sugar Loaf, Co. Wicklow.’ A specimen of the Irish material 
from the Crypt. Coll. Dublin, when examined in 1931, showed very little 
mildew and that only in the Oidium stage. 

5. Gibberella moricola (de Not.) Sacc., Michelia, 1, 347 (1879); Saec, 

ih 553‘ 

On dead fig twigs (jP^rx^i' CanV^x L.), Dorset, January 1946 (W. Buddin). 
Perithecia present in large numbers, mostly immature though a number 
of ripe ones found, free on substratum or partly immersed, scattered or 
aggregated into small tightly packed groups, black (blue in transmitted 
light), 2 10-450 ft (mostly 240-300 ft) in diameter, and mixed here and 

British Fungi. W. C. Moore 87 

there with a few immature buff-coloured ones. Asci cylindrical or club- 
shaped, 60-90 /X long, containing eight monos tichous or subdistichous 
ascospores, sometimes fewer. Ascospores straight or occasionally slightly 
curved, variable in size and shape, usually egg-shaped to spindle-shaped, 
but sometimes broader at one end than the other, 1-5, mostly 3-septate, 
generally with a slight constriction at the septum, 16-25 6-9 [jl (average of 

twenty spores 20x7^), those with two septa broad in relation to their 
length. A few orange-pink sporodochia of a species of Fusarium present 
among the perithecia, with immature sickle-shaped spores, 20-31 x 3-4 ft, 
having acute ends and no discernible septa. 

Wollenweber and Reinking (1935) distinguished Gibberella baccata 
( Walk.) Sacc. var. moricola (de Not.) Wollenw. ( = G. moricola) from G. 
baccata (Walk.) Sacc. by its somewhat smaller spores. Miss E. M. Wakefield 
has, however, drawn my attention to the fact that in SylL Fung, ii, 553, 
Saccardo gives the larger spores (24-30x6-7^6) for moricola and smaller 
spores (16x8 ft) for baccata. But when Miss Wakefield examined two of 
Saccardo’s specimens in Herb. Kewensis, which had been distributed in 
Myc. Ven. nos. 652 and 653 as Botryosphaeria pulicaris f. Mori albae on 
Morus alba^ and B. pulicaris f. baccata on Robinia pseudacacia^ respectively, she 
found that the relative spore sizes were the reverse of those given in the 
Sylloge, The form on Morus had spores which were straight and rather 
cylindric, narrowing only comparatively slightly towards the ends; and 
measuring 16-22 x 5-6 /x. The baccata form had larger, more fusiform 
spores (25-30x6-7/1), most of which were slightly curved. From my 
observations, the material on Ficus from Dorset corresponded more closely 
to the form on Morus than to the baccata form in Saccardo's specimens, and 
it is therefore identified with Gibberella moricola. 

The record of G. moricola on Morus in Hants, listed by Bisby and Mason 
(1940), is an error of transcription: the imperfect stage only of the fungus 
[Fusarium lateritium Nees) was reported in the paper cited. 

6. Leptosphaeria heterospora (de Not.) Niessl, Beitr. 23 (1872); Sacc. SylL 

Fung, II, 67. 

On rhizomes and roots of Iris germanica near Woking, Surrey, March 
1930; on var. Ann Page in St James’s Park, London, October 1930; on 
/. pumila formosa in Kent 1932; and on I. germanica near Aylesbury, 
Buckinghamshire in May 1946. 

The fungus is found on unthrifty and yellowing plants and is probably a 
weak parasite. After making healthy but rather weak growth for several 
weeks in spring, the outer foliage becomes brown and withered, arid the 
younger leaves turn yellow and begin to die back from the tips. On lifting 
affected plants, the rhizomes appear firm and show no decay, but most of 
the roots are withered and dead, or are reduced to discoloured, hollow 
tubes. The black perithecia can be found in appreciable numbers partially 
embedded in the surface of the rhizbmes, especially on the underside, and 
there are usually a few on the roots. The was associated with 

a species of Phoma {with spores 4-6 x i -5-2 />6) in the material from 
Aylesbury, and with Septoria sp. in that from Kent. 

88 Transactions British Mycological Society 

7. Ophiobolus rubellus {Vtrs, ex Fr.) Sacc. Michelia^ ii, 324 (1882) ; Bisby and 
Mason in Trans. Brit myc. Soc. xxiv, 194 (1940); syn. 0. porphyrogonus 
(Tode) Sacc. SylL Fung, n, 338. 

On roots of unpulled and badly weathered flax {Linum usitatissimum L.)^ 
Pembrokeshire (Hayscastle and Dinas Cross) 5 September 1945 (D. L. G. 
Davies) . 

8. Phyllosticta camelliaecola Brun. var. meranensis Bubak, in Ost. bot. Z- 80 
(1905); Sacc. SylL Fung, xviii, 224 (1906). 

On living leaves of Camellia^ St Albans, Herts, May 1939. Spots rounded, 
brown or paling, with a well-defined darker, narrow margin, raised 
especially on the underside, and beheved to be caused primarily by 
insects. Pycnidia on both sides of spots, scattered, small, immersed. The 
spores in this specimen agreed with those of Bubak’s variety (3-5 x i-i -5 jj). 

Phyllosticta camelliaecola Brun. AfiVr. My col. p. 13; Sacc. SylL Fung, x, lox 
(1892), with spores 5-6 x 2-3 /x, is regarded by Grove, Coelomycetes^ i, 10 
(1935), to be the same as P. Camelliae Westd. in Kickx, Flor. Crypt, i, 416, 
for which there are no spore measurements. P. Camelliae was recorded 
from Cound, Salop, in January 1939 (N. G. Preston), but no specimen is 

9. Phomopsis viticola Sacc., in Ann. MycoL^ Berl.^ xm, 118 (1915); Grove, 
Coelomycetes, i, 237 (1935). 

On base of shoot of a vine branch, Dunraven Castle, Glamorgan, May 
1931 (J. Rees). 

10. Spkaeronema pruinosum Peck in Rep. St. Mus. 

On branch of a six-year-old apple tree at Otley (Yorks), April 1928; the 
branch was infested with Mussel Scale and was rather extensively cankered. 

Pycnidia almost superficial, more or less conical or columnar, containing 
macro- and microspores. The larger spores measured 16-27 x 9-1 24^ 
(average of twenty, 23 x loja) and were mixed with large numbers of 
hyaline, straight or slightly curved, unicellular, filiform spores, 6-13 x 1-2 fi 
(average length of twenty, 9*3 />t). The characters of the fungus agreed 
closely with those of Glutinium macrosporum Zeller, described by Zeller (192 7) 
as the cause of a canker of apple and pear trees in Oregon. Examination of 
some of the Oregon material, however, which Dr Zeller sent at my request, 
revealed certain minor differences between the two fungi, which were not 
regarded as sufficient to justify specific distinction. According to Zeller 
(1927) the pycnidia of G. macrosporum produced in culture had smaller 
spores than those (15-28 x 8-1 0-5 /x) found in nature. The spores were 
hyaline and showed only false septation on germination. The fungus in- 
duced a slow rot of apple fruits, but no pycnidia developed on the fruit. 
The spores of the English fungus in culture were up to 30 ^ long. Moreover, 
some of the spores became pale brown, and on germination they sometimes 
became truly i~3-septate. Pycnidia similar to those in culture developed 
on apple fruits slowly rotted by the fungus. 

British Fungu W. C, Moore 89 

Miss E. M. Wakefield, who also examined the two fungi, pointed out 
that G, macrosporum Zeller did not differ from Sphaeromma pruinosum Peck, 
the conidial stage of Pezicula pruinosa Farlow, which was originally described 
on Amelanchier, She examined material in Herb. Kewensis on Amdanchier 
from Farlow’s Herbarium, and also a specimjen collected by J. B. Ellis, 
said to be on elder, which Peck had identified as Sphaeronema pruinosum^ and 
they did not differ from one another or from Glutinium macrosporum. Later, 
Zeller (1935), having examined Peck’s type, accepted this view, though he 
did not regard the fungus as a good species of Sphaeromma, 

11. Ascochyta bohemica Kab. & Bub,, in Hedwigia^ xliv (1905); Sacc. Syll, 
Fung, XXII, 1024 (1913). 

On leaves of Campanula medium L., Harpenden, September 1941 (J. M. 
Gooby). Previously found in Britain on C, betulaefolia and C, Ranieri at 
Maidenhead {Trans, Brit, myc, xxiv, 60). 

On C, medium the spots were round or sub-angular, 2~io mm. in diameter, 
epiphyllous at first, later visible on both sides of leaves, brown, dry, with 
a broad purple margin 2 mm. wide. Pycnidia few, with spores mainly 
unicellular and biguttulate, mostly 12x6 /a, occasionally i -septate, con- 
stricted at the septum and then with several guttules. Perhaps an immature 

12. Ascochyta Cinerariae Tassi, in Boll, Orto, bot, Siena^ 31 (1899); Sacc. SylL 
Fung, XVI, 930. 

On decayed stem bases of cineraria plants in pots, at Carrington and 
Manchester, Lancashire, December 1934 (E. Holmes Smith) ; and at 
Woburn Sands, February 1940 (W. Buddin). 

At Woburn Sands associated with a basal stem rot causing heavy loss 
among plants growing in fresh loam in new pots. Pycnidia numerous, 
aggregated, erumpent, globose or lens-shaped, very variable in size, 180- 
250 /X (-300 jLt) in longer axis (average of ten, 200 /x), pale brown, thin- 
walled parenchymatous, cells somewhat thickened and darker around a 
well-defined ostiole 25-35 p. in diameter. Spores hyaline or slightly 
coloured, egg shaped, 6-10 x 3-5 jcx, mostly i-septate when mature. In 
Lancashire associated with basal stem rot in 40 % of young pot plants. This 
fungus agrees closely with Ascochyta Cinerariae Tassi, which has been 
described as the cause of a leaf spot of cinerarias in Germany (Wasewitz, 
1936) and Italy. The pycnidia are lighter than those of M. {Diplodina) 
Lycopersici^ from which the fungus is otherwise not dissimilar. Diplodina 
(Ascochyta) fibricola (Sacc.), recorded on rotting stems of Cineraria maritima 
in France (Rab. Krypt, FL i^ vi, 682 (1901)), appears to be different. 

It is uncertain whether Ascochyta Cinerariae i% parasitic on the stem bases 
of cinerarias, producing a disease similar to but distinct from Foot Rot 
czMSodi hy Phytophthora cryptogea Pethybr. & Laff., or whether it merely 
develops secondarily on the stem bases of plants primarily affected witlx 
Foot B^ot, ISio Phytophthora was found in the Woburn materiaL 

90 Transactions British Mycological Society 

13. Ascochyta dahliicola (Brun,) Petr,, in Ann, MycoL BerL^ xxv, 201 (1927) ; 
syn, Phyllosticta dahliicola Bran., Champ. Saint. 429 (1887); Sacc. Syll. 
Fung. X, 129 (1892) ; Grove, Coelomycetes^ i, 14 (1935). 

On Dahlia variabilis Desf., Wokingham, Berks, September 1937 (W. 
Buddin) ; var. Clara Carder, Harpenden, Herts, October 1941 (W. Buck). 

14. Ascochyta Impatientis Bres., in Hedwigia^ 326 (1900); Sacc. Syll. Fung. 
XVI, 927. 

On living leaves of Impatiens halsamina L., Newton Abbot, Devon, 
October 1942 (A. Beaumont). 

Pycnidia scattered or aggregated, globose or flattened, immersed, thin- 
walled, pale brown and translucent, 1 20-1 50 fj. diam., opening with a pore 
about 15/X wide. Spores issuing in dense clouds, straight cylindrical with 
rounded ends, 6~ii x 2-4^1, mostly 9 x 3^0., ultimately i-septate, not con- 
stricted at the septum. 

A. Weissiana AJlesch., Rab; Krypt. FI. i, vi, 647 (1899); Sacc. Syll. Fung. 
XVI, 927, on Impatiens halsamina in Saxony, may be the same but is described 
as having black-brown pycnidia, and spores measuring 10-16 x 3'-4*5 ii. 

15. Botryosporium longibrachiatum (Oudem.) Maire, in Ann. My col. Berl.^ i, 
340 (1903) ; Mason in Annotated Account of Fungi received at the Imperial 
Bureau of Mycology, List II (Fascicle i), 27. 

On tomato stems, Rustington, Sussex, September 1943 (W. A. Millard). 

16. Cercosporella Primulae Allesch., in Ber. Bayr. Bot. Ges. ii, 18 (1892); 
Hedwigia, xxiv, 286 (1895); Sacc. Syll. Fung, xi, 607. 

On living leaves of Primula Wanda, Staffs, July 1928; and on hybrids of 
P. Juliae, 'Qxidgt Sellers, Hereford, August 1931, Gamons, Hereford, 
March 1933 (L. Ogilvie), and Bartley, Southampton, May 1936 {Trans. 
Brit. myc. Soc. xxv, 208). 

i^. Fusicladium Lini Sorauer, in Z- TflKrankh. v, 103 (1895); De Wild, & 
Dur. Prodr. FI. 5 ^/^. ii, 336. . 

Forming a black coating on wilted cotyledons and lower leaves of 
usitatissimum L., Egmere, Norfolk (R. E. Taylor) and in Wiltshire (L. 
Ogilvie), June 1942. 

Conidiophores simple, usually fasciculate, septate, pale brown, more or 
less rigid, straight or somewhat bent, ultimately up to over 200 jjl long, and 
3-6 ja broad. Spores borne singly at tips and at times apparently pleuro- 
genous by the continued growth of the conidiophore, mostly straight, or 
sausage-shaped, pale coloured, with a clearly defined wall about i /x thick, 
minutely apiculate, 12-37 x 5-9 /x (average of twenty-five spores 24 x 7/x), 
continuous or i -septate, sub-catenulate. Sorauer did not give a full 
description of his fungus and gave t|ie spores as only 8x4/x, with some 
14-16 /X long. 

Sorauer regarded Fusicladium Lini dis a parasite and the cause of wilting 
and discoloration of the upper portions of flax seedlings in Belgium, 

British Fungi. W. C. Moore 91 

van Poeteren (1929) found what he thought was the same species, or 
possibly only Cladosporium herb arum, on yellow and dwarfed flax seedlings in 
Holland. In England the fungus was associated at both localities with 
yellowing and slow wilting of young seedlings and of plants just beginning 
to flower. 

I am very much indebted to Miss E. M. Wakefield for continued help 
and interest and to all those collaborators who have sent me material. 


Adams, J. (1906). Irish parasitic fungi. Irish Mat. "kvi, 168-9. 

Bisby, G. R. & Mason, E. W. (1940). List of Pyrenomycetes recorded for Britain. Tram. 
Brit. myc. Soc. 197. 

Poeteren, N. van (1929). Verslag over de werkzaamheden van den Plantenziekten- 
kundigen Dienst in het jaar 1928, Versl. PlZi^kt. Dienst Wageningen, no. 58, 15. 
Wasewitz, H. (1936). Schaden durch die Blattfleckenkrankheit der Cinerarien. 

Blwmn- u. PflBau ver. Gartenwelt, XL, 99-100. 

Wollenweber, H. W. & Reinking, O. A. (1935). Die Fusarien. Berlin. 

Zeller, S. M. (1927). A canker of apple and pear trees caused by Glutinium macrosporum 
n.sp. J. agric. Res. xxxiv, 489-96. 

Zeller, S. M. (1935). Some miscellaneous fungi of the Pacific North-west. Mycohgia, 
XXVII, 463, 

{Accepted for publication 12 December 1946) 

[ 92 ] 



Botany Department^ St Andrews University 

Sass (1929) described the agar-film technique by which fungus mycelium 
was grown on microscope slides carrying a thin film of a special semi-solid, 
agar-containing, nutrient medium. Noble (1937) and I (1937) employed 
a modified technique in which the slide carrying the film was placed face 
downwards in a groove cut in a brass box-frame, made to measure to fit the 
slide, and the frame was then put in a Petri dish containing damp blotting 
paper. For examination under the microscope the lid of the Petri dish was 
removed. Light passed to the microscope lenses through the bottom of the 
Petri dish, the film and the slide. The lid could be replaced after examina- 
tion and the fungus allowed to continue growth or, as soon as observations 
of interest were made on nuclear behaviour, the position on the slide was 
marked and noted by mechanical stage readings, the film was fixed, 
stained and examined and the observations made before fixation were 
interpreted in terms of the structures now rendered visible. The worst 
disadvantages of this method are the variable thickness of the Petri-dish 
glass which affects the light, the thickness of the slide and the cloudy 
nature of the medium through which the fungus hyphae have to be 
viewed. Only a very limited number of short lengths of hyphae can be 
brought into focus using a or 3^ in. oil-immersion lens, even when 
‘extra-thin ’ microscope slides are used. 

The following improved technique is now being used in this Department 
in studying the hyphae (particularly the clamp connexions) in certain 
Basidiomycetes. The microscope slide to carry the film is replaced by a 
large glass coverslip 2 X4 in. of the type used to cover fossil preparations, 
or better still by a i x 3 in. no. 3 or no. 2 cover-slip. The bottom of the 
box-frame is set on an ‘extra-thin’ microscope slide. The medium eni- 
ployed for the film is 1*5 % malt in 1*5 % British agar. As before, the films 
are grown in Petri-dish damp chambers. They are removed for examina- 
tion resting on the brass box. The microscope slide completes the base of 
this small damp chamber. Films of this type have been kept growing under 
a in. oil-immersion lens for hours on end — "Sometimes overnight- 
before fixation or return' to their Petri dishes. The small chamb^er is 
comparatively easy to manipulate and hyphae may be traced backwards 
from their tips up to six fields of view under either a 3^ or ^ in, lens. By 
this means the formation of clamp connexions has been watched re- 
peatedly in fungi such Coniophora puteana and Marasmius androsaceus. In 
the latter species the time from the inception of the bulge on the parent 
hypha to the beginning of the outgrowth of a branch from the completed, 

Technique for study of living mycelium. J. A . Macdonald 93 

fused clamp connexion is 50-60 minutes. The formation of the clamp itself 
takes about 20 minutes, with the greatest regularity. 

Marasmius androsaceus forms its clamp connexions in full daylight — 
during ordinary working hours ! — and the technique is simple enough for 
advanced students to be trusted to use the equipment and watch the 
process for themselves. 

An account of results observed is in preparation for publication. 


Macdonald, J. A. (1937). A study of Polyporus betulims (Bull.) Fries. Ann, appl. Biol, 
xxrv (2), 289-310. 

Noble, M. (1937). The raorphology and cytology of Typhula Trifolii Rostr. Ann, BoL, 
Lond,, N.S. I, 67-98. 

Sass, j. E. (1929). The cytological basis for homothallism and heterothallism in the 
Agaricaceae. Amer. J, Bot. xyi, 663-701. 

[Accepted for publication 18 December 1946) 

[ 94 ] 




The Freshwater Biological Laboratory y Wray Castle^ Ambleside^ and 
Department of Botany^ Birkbeck College^ London 

(With Plates IX and X and 8 Text-figures) 

Few workers have been attracted to a detailed study of the aquatic 
Chytridiales in Great Britain (Cook, 1932; Sparrow, 1936; Ingold, 1940, 
1941 and 1944)5 and thus the number of records for this country remains 
relatively small. Intensive work carried out by me on these organisms 
during the past two years shows that they are to be found in almost any 
aquatic habitat provided that a suitable substratum for growth is present. 
Since the Chytridiales are as yet a relatively unexplored group, the 
discovery of new species is not uncommon, and several are described in this 

I. Phlyctidium apophysatum n.sp. 

This chytrid was found on Mougeotia from the Clay Pond, Wray Castle, 
Windermere, in August 1945. Outgrowths from the cells are of common 
occurrence in Mougeotia^ and sporangia of the fungus were often located 
between two such processes. The fungus is parasitic and brings about 
a dissociation of the host chloroplast into numerous granules. 

The thallus is monocentric, and consists of an extramatrical sporangium, 
an intramatrical apophysis and unbranched tubular rhizoid. Young 
stages (Text-fig. la-d) indicate that the apophysis is formed early, as 
a swelling of the germ-tube, the distal part of which develops into the 
tubular rhizoid. The apophysis is spherical to subspherical, and never 
exceeds the diameter of the sporangium. The sporangium is spherical to 
oval, 9-17/X in diameter and usually produces a hundred or more zoo- 
spores, which are liberated upon deliquescence of the apex of the spor- 
angium. The zoospores (Text-fig. i A) are oval, 1-4 x 2-4^6, with a minute 
refractive globule, and a single posterior flagellum about 13/x long; their 
movement is somewhat jerky. The apophysis and sporangium tend to 
collapse after dehiscence. Several smaller sporangia, 7-12 in diameter, 
were observed (Text-fig. le). They contained a few regularly arranged, 
relatively large oil globules, but whether they actually liberate zoospores 
with a conspicuous globule is unknown, as dehiscence was never seen. 
Resting spores were not observed. 

This fungus shows very clearly the dubious nature of the characteristics 
which have been used to separate the genera Phlyctidium and Phlycto- 
chyfhm.l^ resembles a species oi Phlyctidium in the nature of its unbranched 
rhizoid, and a species of Phlyctockytrkm by the possession of an apophysis. 

Studies on British Chytrids. Hilda M. Canter 95 

Sparrow (1933, p. 518, PL 49, fig. 12) described an incompletely known 
fungus on Cladophora^ probably a Phlyctidium^ with a rhizoidal system very 
similar to that of the present species; P. spinulosum (Sparrow^ 19333 p. 5165 
Text-figs, 1 3 2) also has a slightly inflated tubular rhizoid, and it is decided 
to include the fungus here described in the genus Phlyctidium as a new 
species, P. apophysatum^ taking its name from the constant, and well- 
developed intramatrical apophysis. 

Text-fig. I. Phlyctidium apophysatum n.sp, a-d^ young thalli. sporangium possibly containing 
zoospores with a large oil globule, f, dehisced sporangium, g, large sporangium situated 
between two processes of the Mougeotia, A, zoospores, a-f^ h, x 1400; g, x 660. 

Phlyctidium apophysatum n.sp. 

Thallus monocentric, consisting of an extramatrical sporangium 
9-1 7 /X in diameter, an intramatrical apophysis, either spherical 5 ft in 
diameter, or subspherical d-yxiofc, never exceeding diameter of the 
sporangium, continuous with a tubular rhizoid (24 x7 ft) to (12x 3-3 ft). 
Zoospores oval i*4 x 2 *4 ft with a minute colourless globule, and posterior 
flagellum 13 ft long, discharged on deliquescence of the apex of the 
sporangium. Resting spores not observed. 

Parasitic on sp. Clay Pond, Wray Castle, Windermere, 

England, August 1945. 

Phlyctidium apophysatum sp.nov. 

Thallus monocentricus, sporangiis extramatricalibus, apophysibus et 
rhizoideis intramatricalibus. Sporangia g-i 7 ft diam. Apophysis sphaerica 
5 ft diam. aut subsphaerica 6-7-10 ft, nunquam sporangio latior. 

96 . Transactions British Mycological Society 

Rhizoideum tubulare 12 X3*3^ ad 24X Zoosporae ovales, i •4-2-4 
globulo hyalino minuto, postice uiiiflagellatae, flagello 1 3 fx longo, ex apice 
sporangii dissoluto emergentes. Sporae perdurantes non visae. 

Hab. in Mougeotia sp. parasiticum, Clay Pond, Wray Castle, Windermere, 
Anglia, August 1945. 


ScherfFel (1926) described a chytrid parasitic on the sporelings of Oedo- 
gonium which he referred tentatively to Rhizophidium globosum. In February 
1946 a very similar organism was found parasitizing the same host, in 
Clissold Park Lake, London. 

From one to fourteen individuals of the parasite may occur on a single 
sporeling, and when only one is present a characteristic curvature is 
induced in the host cell, the chloroplast of which is converted into a mass of 
brown granules (Text-fig. 2c and PL IX, fig. 4). 

The rhizoidal system is often difficult to observe owing to the dense 
chloroplast of the host, but where visible it consists of a main axis, rarely 
slightly swollen immediately beneath the host wall (Text-fig. 2 a, b)y which 
branches to give a meagre rhizoidal system. The spherical sporangia vary 
in size from 7-5 to 34*3 /x in diameter, and where many occur on a single 
host cell they are relatively small. 

As the sporangium matures a single lateral dehiscence papilla becomes 
visible, and it appears to be filled with a plug of highly refractive material. 
The part of the sporangium wall forming the papilla deliquesces, but 
whether the plug also deliquesces or is extruded as a solid mass is unknown, 
as the actual moment of dehiscence was never observed. Hundreds of 
posteriorly uniflagellate zoospores 4 (jl long, 2 /x diameter, are produced. 
They are of unusual structure for a chytrid since they have no conspicuous 
oil globule, but they contain one or two minute, highly refractive granules, 
often situated laterally near the posterior end (Text-fig. 2^). 

Spherical, asexually formed resting spores 21-4-35^6 in diameter were 
seen, which appeared to produce little effect on the host chloroplast 
(Text-fig, 2h and PI. IX, fig. 6). They are similar to the zoosporangia, but 
the wall is up to 2 p thick; their contents are at first granular, but later 
become oily. Neither the rhizoids nor germination of these resting spords 
was seen; one empty specimen showed a single lateral dehiscence pore. 

The chytrid here described agrees in all essentials, except one with 
\ScherfFers organism. In the latter the rhizoidal system is stated to consist 
of a few fairly thick but not long, forked rhizoids, arising from the base of 
the sporangium (Text-fig. 3), whereas in the London material there is 
a single main axis. In my opinion this is not a significant enough difference 
to warrant a separation of the two forms. 

Although the chytrid here described is very similar to R, globosum^ 
according to Sparrow’s (1943) definition it cannot find its true place in this 
species owing to the smooth-walled resting spore. Great difficulties are 
presented in dealing with the globose species of Rhizophidium, Tht records 
are numerous, but only rarely is a complete description given. Observa- 
tions on the structure and method of formation of the resting spores is of 

Text-fig. 2. Rhizophidium sp. a, very yoiing sporangia; intramatrical rhizoid is slightly swollen 
immediately within the host wall, c, sporangium with well-developed rhizoidal system. 
d, immature sporangium rhizoids not visible; characteristic curvature of the host present, 
but contents little disorganized, e, mature and dehisced sporangia. mature sporangium 
with lateral plug of highly refractive material, g, zoospores, smooth walled resting spore. 

X 975; a, X 1333; r, X 700; fif, X 700. 





98 Transactions British Mycological Society 

utmost importance^ and when investigations have been carried out on the 
specificity and morphological variations of these organisms, a thorough 
revision of the genus will be necessary, 

III. Rhizidium variabile n.sp. 

This fungus was found growing as a saprophyte on Spirogpm in May 1945 
and 1946, from a pond in Chelsea Physic Garden, London. It appeared 
after the Spirog^ra had been kept in the laboratory for some weeks, by 
which time the algal cells were almost unrecognizable. 

The thallus is monocentric, consisting of an extramatrical, usually un- 
branched system of diverse form and size, and an intramatrical ultimately 
branched rhizoidal system. The extramatrical part is composed of the 
body of the encysted zoospore, more or less swollen, continuous with 
a single stout rhizoid, which may exhibit one or several swellings through- 
out its length. The zoospore on germination produces a germ tube which 
branches distally, the swellings developing later. The diversity of the thallus 
structure is illustrated in Text-fig. 4 5 and PL X, figs. 4-6. The exact point 
where the thallus enters the host cell is often difficult to determine, because 
of the disorganization of the cell walls. In the material found in 1945 
(Text-fig. 4;C) the intramatrical system of the chytrid was rarely swollen, 
whereas in the collection made a year later many such specimens were seen 
(Text-fig. which simulated species of the genus Phlyctochytrium. 

During the development of the vegetative part of the thallus the encysted 
zoospore remains small, later there accumulates in it the protoplasmic 
contents of the whole thallus, and it enlarges to form a spherical or sub- 
spherical sporangium, 6-13 in diameter, which contains from one to 
twelve relatively large zoospores. These exude in a motionless mass on 
deliquescence of the apex of the sporangium. At first they are amoeboid, 
but soon round off and swim away. The zoospores are 4-4-5 fx in diameter 
with an oil globule 1-2 [x in diameter and a single posterior flagellum 26 /x 
long. The sporangium wall collapses after dehiscence. Resting spores were 
not observed. 

In general structure this species most nearly rtsemhlts Rhizidium 
mycophilum Braun, previously recorded from England by Sparrow (1936), 
growing on exuviae of Ghironomidae. However, the series of subsporangial 
swellings is not a feature of R. mycophilum^ and the sporangia and number 
of zoospores produced are much smaller. Further, the zoospore mass does 
not exhibit the changes in shape that were recorded for Sparrow’s material 
of i?. mycophilum. Karling (1944) recognizes i 2 . Braun as two 

species, iJ. Braun and R. Mowakowskii Karling ( ==i?, mycophilum 

Nowak), based on differences in habitat, in the structure of the zoospores, 
and in the resting sporangia. As neither Sparrow (1936) nor I have 
observed resting spores the exact affinity of the present species cannot be 

It is suggested that the present fungus sha]l be described as a new species, 
until experiments have been carried out on the morphological variations 
which occur in single-spore cultures on different hosts in R. variabile or its 

100 Transactions British Mycological Society 

near allies, after which it may be necessary to revise the position of this 

Rhizidium variabile n.sp. 

Thallus monocentric, extramatrical part consisting of a sporangium 
6-13 p. in diameter (developed from the encysted zoospore), and a single 
stout rhizoid, which may have one or more swellings. Intramatrical part 
sometimes swollen, tapering to a branched rhizoidal system. Sporangium 
containing 1-20 zoospores which emerge, surrounded by a vesicle, on 
deliquescence of the sporangial apex. Zoospores spherical 4-4-5 p in 
diameter, with a single oil globule r-2 p in diameter, and a posterior 
flagellum 26 p long. Sporangium wall collapsing after dehiscence. Resting 
spores not observed. 

On dead Spirogyra sp., Chelsea Physic Garden, London, England. 

Text-fig. 5. Rhizidium variabile n.sp. a, b, young thalli with slender branched intramatrical 
rhizoidal system, c, thallus with extramatrical and intramatrical apophyses, d, empty 
sporangium, above two zoospores, e, empty collapsed sporangium. (Chelsea Physic Garden, 

Rhizidium variabile sp-nov. 

Thallus monocentricus. Pars extramatricalis e sporangio et rhizoideo 
unico dilatationibus singulis vel pluribus praedito consistens. Pars intra- 
matricalis interdum inflata, in rhizomycelium ramosum attenuata. 
Sporangium 6-13 p diaroetro, zoosporas 1-20 includens. Zoosporae 
sphaericae, 4-4-5 p diam., globulo unico hyalino i-2 p, postice uniflagel- 
latae, flagello 26p longo, vesiculo inclusae emergentes. Membrana 
sporangii post dehiscentiam collabit. Spore perdurantes non visae. 

Hab. in Spirogyra sp. emortu'a, Chelsea Physic Garden, London, Anglia. 

Studies on British Chytrids. Hilda M. Canter loi 


This chytrid (Text-fig. 7, and PL X, fig. 7, 8) was found growing on 
JSfitzschia sigmoidea (Ehrenb.) W.Sm., in Bradbourne Park Lake^ Sevenoaks, 

Kent, England, from November 1944 to March 
1945. It was also found in a collection from the 
River Yeo, Sherborne, Dorset, in March 1945. 

Although other diatoms were abundant from 
both localities (e.g. species of Pinmlaria^ Syne-- 
dra^ Mavicula^ Surirella^ Melosira and Gyrosigma)^ 
they were never attacked. It therefore seems 
probable that the chytrid is specific to Mitzschia 
sigmoidea^ as there should Ixave been ample 
opportunity for infection to occur in the small 
flocculent masses into which these diatoms 
were crowded. The fungus is parasitic, but the 
contents of the infested cells are little affected ; 
the chromatophore retracts somewhat at the 
point of entry of the rhizoids, but the movement 
of the host is not impaired. An infected diatom 
may carry from one to thirty individuals of the 
parasite (Text-fig. 6). 

The zoospore encysts on the surface of the 
diatom and produces a germ tube which usually 
penetrates a carinal dot. From the base of the 
germ tube a branched intramatrical rhizoidal 
system arises (Text-fig. 7^). The mature spor- 
angia are obpyriform and taper to a knob-like 
base sessile on the host. The sporangium wall 
is smooth, colourless, thin distally, often be- 
coming thickened at the base. The sporangia 
range from 4 to 25JL6 in diameter, and from 
15 to 60 p, high; while the smaller sporangia 
contain a few zoospores, the larger ones may 
produce eighty or more. The zoospores are 
spherical, 3-4 p, in diameter, uniguttulate with 
a single posterior flagellum, and upon the de- 
tachment of an apical, rarely somewhat lateral 
operculum, they escape in a mass from the 
sporangium but quickly separate, and glide 
away individually. One probable resting spore 
was observed (Text-fig. 7 A), but it is thought 
unwise to base a description on this specimen 

Apart from the absence of a stalk, its larger 

size and supposed specificity for Mitzschia sig- Text-fig, 6. Chytridium versatile vzx, 
moideay the fungus agrees well with Chytridium MtzsMas^i^rnoideah^Qxmg 

versaMle Scherffel, already recorded from this ous^fagesof development^ 

102 Transactions British Mycological Society 

country by Sparrow (1936). As in C. versatile (see ScherfFel, 1926) the 
sporangium bends back as the diatom pushes against debris in its en- 
vironment, in spite of the fact that this species is sessile. After the 
obstruction is passed the sporangium snaps back to its original upright 
position. ScherfFel (1926, Plate 9, figs. 19, 20) figures three sporangia 
apparently without a stalk; however, no mention is made in the text to 
such sessile forms. Owing to the absence of any major structural 
difierences from C. versatile ScherfFel it is proposed to erect a new variety, 
C. versatile var. acaulis, being characterized by the absence of a stalk. 

Text-fig, 7. Chytridium versatile var. acaulis. a, young sporangia, b, sporangium inclined to long 
axis of host cell, c, mature sporangium, d, dehisced sporangium showing branched rhizoidal 
system, e, zoospores. /, g, dehisced large and small sporangia, resting spore (?), di, 
X 1400J a, by e,fy g, hy x 660. 

V. Chytridium cocconeidis n.sp. 

This fungus was found on Cocconeis pediculus Ehrenb., itself epiphytic on 
Cladophora, in a small lake in Clissold Park, London, March 1945. Sparrow 
(1943) records no chytrid on Cocconeis, and the organism appears to be an 
undescribed species. ' 

Judging by the disorganized state of the chromatophores in the affected 
diatoms, the chytrid is parasitic. It seems to be limited in its host range, 
since Rhoicosphenia curvata (Kg.) Grun., which was also abundant on the 
was not attacked. 

The species is monocentric, and the thallus consists of a cylindrical 
sporangium, and a rhizoidal system. This system consists of an extra- 
matrical part, and presumably also of an intramatrical part, although this 
was not demonstrated microscopically. The extramatrical part consists 
sometimes of a straight imbranched portion (Text-fig. 8 a, g, i), but at 

104 Transactions British Mycological Society 

other times it is forked near the attachment to the host cell (Text-fig. 
8 ^5 e) . Occasionally no extramatrical part was visible, and the sporangium 
appeared to be sessile on the Cocconeis (Text-fig. 8r, i). Mature sporangia 
measure 1 5--29^ in length, by 5-6^6 in diameter, and are sometimes 
inclined at an angle to the main extramatrical rhizoid. The sporangium 
contains from twelve to thirty zoospores and dehisces by the separation of 
a convex apical lid 2-3 /x in diameter. Upon detachment of the lid several 
zoospores escape together, but the remainder emerge singly. The zoospores 
are spherical, 2-3 /x in diameter, with a conspicuous refractive globule and 
a single posterior flagellum. Their movement is predominantly hopping, 
with periods of gliding. The method of infection of the Cocconeis by the 
zoospore was not satisfactorily demonstrated. Resting spores were not 

Since this chytrid is operculate, in Sparrow’s classification (1943) it 
belongs to the Chytridiaceae. The only possible genus appears to be 
ChytridiuiUy but this genus contains no species with a branched extra- 
matrical rhizoidal system, although a few species, C. versatile Scherffel, 
C. curvatum Sparrow, and C. Lagenula Braun pro parte, may have a short 
slender extramatrical stalk. 

Although the inclusion of this species in the genus Chytridium may 
necessitate a slight extension of the concept of the genus, this is nevertheless 
preferable to the erection of a new genus. The species C. cocconeidis is 
proposed, taking its name from the host upon which it is apparently a 
specialized parasite. 

Chytridium cocconeidis n.sp. 

Thallus monocentric, eucarpic, sporangium extramatrical cylindrical 
15-29 in length, 5-6 [m in diameter, dehiscing by an apical lid, and con- 
taining 12-30 zoospores. Zoospores spherical 2-3^ in diameter, with a 
colourless globule and single posterior flagellum. Extramatrical rhizoidal 
system simple or branched 2-9 ju in length, rarely absent. Intraihatrical 
rhizoidal system not observed. Resting spores not observed. 

On living cells of Cocconeis pediculus from Clissold Park, London, England, 
March 1945. 

Chytridium cocconeidis sp.nov. / 

Thallus monocentricus, eucarpicus. Sporangia extramatricalia, cylind- 
rica, 15-29x 5-6^, operculo apicale dehiscentia, 12-30 zoosporas in- 
cludentia. Zoosporae sphaericae 2-3 /x diam., globulo hyalino refractivo, 
postice uniflagellatae. Rhizoidea extramatricalia, simplicia vel ramosa, 
2-9 /X longa, raro nulla. Sporae perdurantes non visae. 

Hab. in cellulis vivis Cocconeidis pediculi, Clissold Park, London, Anglia, 
Martio 1945, 


Five monocentric chytrids are described growing on algae from Great 
Britain. Of these, three are new species, namely, apophysatum, 

Rhizidium variahile Chytridium cocconei£^^ Chytridium versatile var. acaulis 

Studies on British Chytrids. Hilda M, Canter 105 

represents a new variety, and a species of Rhizophidium is identified with 
Rhizophidium globosum Schtrffel {ig26) . 

My thanks are due to Miss E. M. Wakefield for the Latin diagnoses and 
to Prof. C. T. Ingold for the advice he has given throughout the course of 
this work. 


Cook, W. R. I. (1932). An account of some uncommon British species of the Ghytridiales 
found in algae. Mw PhytoL xxxr, 133-44. 

Ingold, G, T. (1940). Endocoenobium Eudorinae, gen. et sp.nov., a chytridiaceous fungus 
parasitizing Eudorina elegans Ehrenb. Mew PhytoL xxxix, 97-103. 

Ingold, G. T. (1941), Studies on British Ghytrids. I. Phlyctochytrium proliferum sp.nov. 
and Rhizophidium Lecythii sp.nov. Trans. Brit, mycol. Soc. xxv, 41-8. 

Ingold, G. T. (1944). Studies on British Chytrids. II. A new Ghytrid on Ceratium and 
Peridinium. Trans. Brit, mycol. Soc. xxvii, 93-6. 

ICarling, J. S. (1944). Brazilian chytrids II. New species of Rhizidium. A. J. Bot. xxxi, 

ScHERFFEL, A. (1926). Einiges xiber neue oder ungeniigend bekannte Chytridineen 
(Der ‘Beitr%e zur Kenntnis der Chytridineen’, Teil II). Arch. Protistenk. liv, 167- 

Sparrow, F. K. (1933). Inoperculate chytridiaceous organisms collected in the vicinity 
of Ithaca, N.Y., with notes on other aquatic fungi. Mycologia, xxv, 513-35. 

Sparrow, F. K, (1936). A contribution to our knowledge of the aquatic Phycomycetes of 
Great Britain. J. Linn. Soc. Lond. {Bot.)^ l, 417-78. 

Sparrow, F. K. (1943). Aquatic Phycomycetes. Ann Arbor, U.S.A. : University of Michigan 

Plate IX 
Rhizophidium sp. 

Fig. I . Many germinated zoospores on an Oedogonium sporeling. x 700. 

Fig. 2. Two sporelings each with a young sporangium; the one on the right already shows 
a slight curvature, x 650. 

Fig. 3. Sporeling attacked by five sporangia, the chloroplast is very much contracted. 

Fig. 4. Almost mature sporangium, x 650. 

Fig. 5. Sporangium showing lateral region of dehiscence, the outer wall has already deliquesced, 
X 650. 

Fig. 6. Resting spore with thick wall; the host contents are not disorganized. X650. 

Plate X 

Fig. I. dehisced sporangium with zoospores at its apex. X 480, 

Fig. 2. Sporangium with needle-like extramatrical rhizoid. X 640. 

Fig. 3. Cylindrical, apparently sessile sporangium on Cocconeis pediculus. x 960. 

Fig. 4. Rhizidium variabile; young thallus showing encysted zoospore, intramatrical apophysis and 
rhizoidal system. X 560. 

Fig. 5. Thalli visible at {x,y, ^), (x) shows a long extra matrical rhizoid; the cell walls of the 
are just out of foc^^^ X 280. 

Fig. 6. At (a) is a mature sporangium with the oil globules of six zoospores clearly delimited; 
a young thallus is visible to the right of (a) . X736. 

Fig. 7. Chytridium versatile var. acaulis^ a mature sporangium on Mitzschia sigmoidea. x 700. 

Fig. 8. Two dehisced sporangia, x 190. 

[Accepted for publication 20 December 1946) 

[ io6 J 


By W. P. K. FINDLAY, D.Sa 

Department of Scientific and Industrial Research Forest Products Research Laboratory 

(With 2 Text-figures) 


The use of perforated cards for the rapid identification or selection of 
individuals possessing any particular combination of characteristics is now 
well known. The method has been successfully used for the identification of 
timbers from their microscopic structure and other features (Clarke, 1938; 
Phillips, 1941). An attempt has now been made to construct a perforated 
card key for the rapid preliminary identification of Hymenomycetes 
belonging to Polyporaceae and Agaricaceae. 

Many would-be students of these groups are discouraged by the initial 
difficulties which they experience when they attempt to identify specimens 
by means of the orthodox methods, such as dichotomous keys based on a 
few characters which may appear to them ill-defined. Nothing encourages 
students more than to make by themselves a number of successful determina- 
tions, and it is with the hope of encouraging them to take an interest in the 
higher Basidiomycetes that the following suggestions are published. 


Cards with small marginal perforations are marked out with a list of the 
most useful diagnostic features. Cards printed with a list of these features 
may be prepared* or the features may be listed only on a master card and 
corresponding blank cards prepared for each species to be included in the 
key. The cards are prepared for use by clipping out the edge of the holes 
opposite to the number assigned to these particular features that are 
possessed by the species in question ; for instance, if a species is white, soft 
and possesses a fixed ring on the stem the holes assigned to these characters 
are clipped out. All the cards of the group that is under consideration are 
stacked together in a box, the right way up and facing the same wayyOnt clipped 
corner of the card providing a check that this is done. It is then immaterial 
how they are arranged in the ^ pack’. 

When using the key the unknown specimen is examined and the most 
important diagnostic feature is noted. A knitting needle, preferably 
mounted in a wooden handle, is then run through the hole through the 

* The right to produce cards of this kind is covered by patent held by Messrs Copeland 
Chatterson and Go., London, E.C. 4, 

Cards for Identification of FungL W, P, K. Findlay 107 

entire pack of cards, which is gently but thoroughly shaken. All the cards 
describing species which possess this feature fall out ; a second well-marked 
diagnostic feature is then chosen and the needle passed through the corre- 
sponding hole in the cards which have fallen out, the process being continued 
until one card or only a few cards are left. If only a few cards are left it 
is then usually possible to select the card descriptive of the unknown 
specimen, since additional notes on characteristic features that are not 
included in the standard characters assigned to the perforations are written 
on the cards to assist final diagnosis. 

The successful working of any key depends, of course, on the choice of 
well-defined, constant characters on which to base it. 

It is recognized that one of the principal difficulties in satisfactorily 
describing fungi for purposes of subsequent recognition is the paucity of 
well-marked characters — shape, size and colour often being so variable as 
to afford little assistance, whilst other more constant characters, such as 
texture, are difficult to describe precisely. Such difficulties arise in the 
construction of any type of diagnostic key, but the perforated card system 
provides a loophole by which variations can be covered, e.g. if a species is 
black or white, according to conditions under which it is grown, both of the 
holes corresponding to these characters are clipped so that the card for this 
species falls out whether the specimen belongs to the black or the white 
variety, and its card will be among those chosen for further selection. 

Advantages of perforated card system 

(1) Any striking characteristic may be chosen to make the first selection, 
whereas with a dichotomous key the choice of characters and the sequence 
in which they are made must follow the lines laid down by the author of the 
key. A few examples will make this clear. In practice one often recognizes 
a species by its odour or by its change of colour on wounding — such 
features cannot obviously be made the basis of a formal dichotomous key, 
but using the perforated cards one can at once select those species having 

such striking characteristics. 

(2) Card key systems permit the inclusion of species with features 
untypical of the genera to which they belong which have to be treated as 
exceptions in dichotomous keys: for instance, there are species that possess 
coloured spores that are normally included in general ]ik.^ Lactarius and 

which in some classifications (e.g. Ramsbottom, 1923) are described 
for the sake of simplicity as possessing white spores. 

(3) New species can readily be introduced into the key without affecting 
the position of those already included. 

(4) The number of species included may amount to many hundred 
without rendering the key unworkable. 

While the number of alternative characters that can be included in 
a card key cannot conveniently exceed 1 20-1 30, the blank space in the 
centre of the card can be used on which to write supplementary informa- 

io8 Transactions British Mycological Society 

The general keys to the Polyporaceae and Agaricaceae illustrated below 
are intended to serve as guides only to well-marked species or to groups of 
species. In some genera they will obviously serve only to indicate the 
groups of species that some taxonomists will regard as subgenera or genera. 
I wish to emphasize that it would not usually be possible, for instance, to 
run down a species of Cortinarius or Russula from a general key to the 
Agaricaceae of any region, but the key should lead to the group of the 
particular genus, which can then be studied in detail from a recognized 
flora. It should be possible for anyone interested in one of these difficult 
groups to construct for themselves a card key restricted to that particular 
genus in which the characteristics of the species in the genus are given in 
greater detail than is possible in the general key. For instance, the 
diagnostic characters of the species of Boletus as described by Pearson 
(1946) could well be used as the basis on which to construct a perforated 
card key. 

Use of perforated card key 

It is obviously necessary for the user of a card key to be familiar with the 
features listed and to have some idea of their relative importance for 
diagnostic purposes. In using the key for Agaricaceae one would naturally 
choose first such features as spore-colour, attachment of gills, texture of 
pileus and stem, and so on. The characters printed on the card are the main 
basic characters, those assigned to the holes lettered <2, r, etc., are those 
which amplify or extend the range of the main characters. On the 
Agaricaceae card, a-f^ for instance, are additional colours, g-l give details 
of gills, m-r of spores and hymenium, while s-y describe the upper surface 
of the pileus in greater detail. 

When using the key it is important to realize that some of the features are 
to be understood as relative to the size of the fungus, for instance, flesh 
thin’ or ‘flesh thick’ is not an absolute quantity in the way that the overall 
size is — such features should be used only after the more definite characters 
have been employed. 

Fig. I shows a sample card from the card key to the Agaricaceae clipped 
to d^scvihc Amanita muscaria\ for this readily recognized species three 
features are sufficient to sort out this particular card from the pack, (i) the 
volva, (2) the fixed ring, (3) the colour of the pileus. 

Subsidiary characters not printed on the card for Agaricaceae : 

a. Pileus fuscous m. Spores pip-shaped 

h. Pileus ferruginous n. Spores curved-allantoid 

c. Pileus yellow-brown 0. Spores apiculate 

Pileus olivaceous p. Spores guttulate 

e. Pileus fawn or bistre Spores amyloid 

/. Pileus pallid or off-white r. Cystidia present 

g. Gills adnexed s. Upper surface of pileus tomentose 

h. Gills emarginate U Upper surface of pileus pruinose 

L Gps venyicose u. Upper surface silky or fibrillose 

j. Gills havmg irregular edges v. Flesh blueing on wounding 

k. Gills having crisped edges w. Flesh reddening on wounding 

L Gills anastomosing or connected by veins x Pileus umbilicate 

j. Pileus repand 

Cards for Identification of Fungi. W. P. K. Findlay 


c 0 0 0 0 0 


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06£ WNOWW «0 ONOtJiS 




QlbOV ** 

OllW 31SV1 


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Transactions British Mycological Society 

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Cards for Identification of Fungi. W. P. K. Findlay ill 

Fig. 2 shows the characters included in the key to the Polyporaceae. 

Subsidiary characters not printed on key for Polyporaceae: 

a. Upper surface fawn d. Upper surface sulcate 

Upper surface ferruginous e. Upper surface sticky 

c. Upper surface pallid 

The characters on which to base the key have been chosen as being those 
which can generally be determined with minimum delay^ and the 
characteristics of the fungi when grown in culture on agar medium have 
not been included although in the case of some genera of Polyporaceae, e.g. 
Poria spp., they are very helpful. A similar card key for the identification of 
cultures of wood-rotting fungi is now in the course of preparation and will 
be published later if it proves useful. Only by using these keys will their 
flaws and omissions become apparent, and I shall be grateful for any 
suggestions or criticisms that occur to other workers. 


A method is described for the preliminary identification of Agaricaceae 
and Polyporaceae by the use of perforated cards. Specimen cards listing 
the features used for the diagnosis of species in these two families are 

This work has been carried out as part of the programme of the Forest 
Products Research Boards of the Department of Scientific and Industrial 
Research, by whose permission it is published. 


Clarke, S. H. (1938). A multiple-entry perforated-card key with special reference to the 
identification of hardwoods. New PhytoL cxxxvii, 370--4. 

Pearson, A. A. (1946). Notes on the Boleti. July-September. ^ 

Phillips, E. W. J. (1941). The identification of coniferous woods by their microscopic 
structure. J, Linn* Soc* Lond. lix, 259-320. 

Ramsbqttom, J. (1923). A Handbook of the larger British Fungi, London. 

[Accepted for publication 20 December 

[ II2 ] 


By H. wild, B.Sc., Ph.D. 

From the Department of Botany, Imperial College of Science and 
Technology, South Kensington, London 

A. Introduction 

Mildew of lettuce, caused by Bremia Lactucae Regel, is a disease of long 
standing and one which occurs more or less in all places where lettuce is 
grown. It has been reported from all continents and from both temperate 
and tropical regions. The disease is readily recognized and is not likely to 
be confused with any others which infect the crop, and there has never 
been any doubt as to the identity of the causal organism. While, therefore, 
so much is definite, it is somewhat surprising that certain features of the 
disease are still largely not understood. Thus it is not clear by what means 
the fungus carries over from one crop to a succeeding one — ^in particular, 
it is not known whether the disease is transmitted by means of contaminated 
or infected seed or by infected residues of earlier crops. Morphologically 
the same species occurs as a parasite on various composite weeds (groundsel, 
Sonchus, etc.), and it is not certain how far these may function as inter- 
mediate hosts. 

Again, though the disease is nearly always to be found without difficulty 
upon growing crops, there is little substantial evidence bearing on the 
damage which it produces. Affected plants, as a rule, are not killed, and 
they may in the long run mature satisfactorily. In the language of the 
cultivator, the plant often 'grows away from the disease’, by which is 
meant that it does not die, but finally produces a saleable lettuce. It does 
not follow, however, that the effect of the disease is negligible — maturity 
may be delayed or the size of the mature plant may be reduced. Clear 
evidence on this point is lacking. 

The work described in this paper was carried out over the two seasons 
1939-40 and 1940-1 and relates mainly to the points raised in the fore- 
going paragraphs, viz. the mode of transmission of the disease, and the 
influence of the disease on the maturing of the crop. Fungicidal methods of 
control have also been examined. These problems have been studied under 
the conditions of cultivation which prevail in the market-garden area which 
lies in the Thames valley west of London, and as the mechanism of disease 
transmission is obviously related to the methods of cultivation practised, 
a short account of the latter is desirable. 

B. Methods of cultivation 

Beginning in September, winter lettuce is sown in the open, followed by 
a later sowing in frames from late October to early November, the latter 
crop being planted out in spring. These crops mature from about early 

Downy Mildew Disease of Cultivated Lettuce. E. Wild 113 

May to the middle of June. During approximately the same period, i.e. 
from September to April, lettuce crops may be grown and matured in 
frames (cold or heated) or as catch crops in greenhouses. Finally, summer 
lettuce is sown in the open from March onwards, and these crops mature 
from July to as late as December, if the early winter is not too severe. 

Thus on any market garden in the area round London, there is much 
overlapping of lettuce crops. It is particularly noteworthy that winter 
lettuce, which constitutes a fair proportion of the new year’s early field 
crop, overlaps by about two months the last of the preceding year’s summer 
crop. The weather in October— December is often very favourable to 
mildew, and in fact this disease is often rampant in late summer crops. The 
new crop cannot always be segregated from such infected earlier crops and 
therefore there is the possibility of wind-borne spores pasring from the one 
to the other. 

My attention was directed chiefly to the incidence of mildew in ' Spring 
lettuce’, i.e. lettuce which is sown in unheated frames in late October or 
early November and planted out from February to April. In such frames 
one frequently sees isolated seedlings with one or both cotyledons carrying 
mildew. These occur sporadically, though later whole areas of the frames 
may become affected. The first outbreak of the disease thus appears to 
suggest seed-borne transmission. 

C. Transmission of mildew by seed 

The method used for investigating the possibility of mildew being seed- 
borne was as follows. Seed-boxes were dipped for five to ten minutes in 
boiling water and filled with soil which had been autoclaved for one and 
a half hours. In order to safeguard against air-borne contamination, the 
boxes after being seeded were covered with glass plates or bell-jars. In 
later experiments, an improved device, viz. a large glass-sided frame with 
closely fitting glass lid, was used. The boxes were freely watered with 
sterile water, and thus a highly humid atmosphere, favourable to the 
development of mildew, was provided. 

From three to six weeks after sowing, according to the time of year, the 
seedlings had developed their first two foliage leaves. It could thus be 
assumed that any seed-borne infection would have had time to produce 
conidia on the cotyledons, and so the seedlings were removed singly with 
forceps and examined for the presence of mildew. 

In the first five experiments of this kind Trocadero Improved, Imperial 
and Early French Frame seed of the current year (1939) was used. Later 
(Exps. VI-VIII) older seed of Trocadero was tested as the 1939 stock may 
have been unusually clean. The commercial seed so far used may, however, 
have been harvested from plants which were uninfected by mildew and so 
in the autumn of 1940 seed was harvested from Imperial, Trocadero, and 
Winter King plants, the first of which were known to have been heavily 
infected, but the other two had only been slightly infected and then not at 
the harvesting period. With the expectation that seed of Imperial mother 
plants so infected would be most likely to show whether seed-borne 



1 14 Transactions British My cological Society 

transmission of mildew does or does not occur, four experiments (XI-XIV) 
were carried out with this seed at various times over the winter of 1 940-1. 

The data obtained from the fourteen experiments bearing on the 
question of seed-borne transmission are assembled in Table i. The results 
may be summarized as follows : 

(1) Imperial, Trocadero and Early French Frame seed of 1939 showed no 
seed-borne transmission of the disease. 

(2) One sowing of commercial Trocadero seed (Exp. VI) contained 
a small percentage of infected seedlings but the other two did not. 

(3) Two experiments (XI and XII) with seed taken from infected mother 
plants gave slight infection but four did not. 

(4) The total number of seedlings examined from October 1939 to May 
1941 was 23,755 and of these only 0*025 % were infected with mildew. 

Table i. The percentage seed infection of lettuce by mildew 


Seed sown 





% infected 



i. x. 39 




1939 Troc. 


15. xii. 39 




1939 Imp. 


15. xii. 39 




1939 E.F.F. 


15. xii. 39 




1939 Troc. 


25. i. 40 




1939 IniP- 


29. iii. 40 




1938 Troc. 


29. iii. 40 




1938 Troc. 


20. V. 40 




1936 Troc. 


24. ix. 40 




1940 W. King 


24. ix. 40 




1940 Troc. 


24. ix. 40 




1940 Imp. 


23. X. 40 




1940 Imp. 


1 1 . xii. 40 




1940 Imp. 


10. iii. 41 




1940 Imp. 

D. Transmission by debris of infected crops 

Lettuce frames are often set up in the same place as in the preceding years, 
and succeeding field crops sown in the same ground, because market 
gardeners in this area use intensive methods of cultivation and cannot 
practise a long-term rotation system. It is possible therefore that viable 
fungal residues may carry the disease over in the field or in frames, 
particularly in the latter. This brings into consideration the question of 
resting structures, in particular oospores, and of the viability of mildew 

Oospores of Bremia Lactucae were first described by de Bary (1863) but 
from Senecio vulgaris only. References to their occurrence on the cultivated 
lettuce are given by Smith (1884), Schweizer (1919), Lavrov (1932), and 
Baudys (1935), and from these it is apparent that oospores are rarely found 
on that host. 

An attempt was made to find them in the debris of lettuce plants which 
were heavily infected with mildew when alive. Lettuce leaves have a high- 
water content and a low proportion of lignified tissue, so that they rapidly 
disintegrate and disappear soon after falling to the ground. If, however, the 
soil is not too wet they become dry and after a few months in winter are 

Dozony Mildew Disease of Cultivated Lettuce. H. Wild 115 

black and friable. For microscopic examination this type of material was 
cleared by immersion in a mixture of equal parts of chloral hydrate and 
pure phenol crystals. These mix on being gently warmed, and on cooling 
the mixture remains hquid. The fragment of leaf was then placed in 10 % 
caustic potash solution for a few minutes, washed in several changes of 
water, stained with aniline blue and finally cleared with lactic acid. 

Many slides were prepared in this way from decayed lettuce leaves, but 
no oospores were seen. On the other hand, recognizable mycelium of 
BremiaLactucae was almost invariably present, and it is possible that portions 
of this mycehum may long remain alive. 

To test whether soil containing debris from infected plants could infect 
seedlings, a stock of soil was built up between December 1939 and May 
1941 by the continued addition of badly infected leaves to a large box of 
soil. At intervals some of this soil was removed and about one-third of the 
sample autoclaved at 15 Ib./ for hours to be used as control. 
Commercial seed was then sown in both lots of soil and protected from 
wind-borne spores as in the experiments described in the previous section. 
The varieties used were Trocadero and Imperial, the earlier tests having 
shown that the commercial seed of these varieties did not carry mildew. 
To encourage development of mildew the humidity was kept at a high 
level by watering freely with sterile water. After three or four weeks the 
seedlings were examined for infection, with the results shown in Table 2. 

Thus, in all, over 5000 seedlings were grown under conditions highly 
favourable to the development of mildew, but only one infected seedling 
was seen. As much debris was present it seems that any viable fungal 
remains in it were ineffective in transmitting the disease. 

Table 2. Effect of contaminated soil on the occurrence of 
mildew in lettuce seedlings 
Contaminated soil Controls 


Date of sowing 

No. of 




No. of 





13. iii. 40 







13. iii. 40 







3. V. 40 







8. V. 40 







22. X. 40 







g.iv. 41 











The possibility of a carry-over by conidia is unlikely but it was desirable 
to determine the time limit of their viability. For this purpose a number of 
slides were plentifully dusted with conidia, placed in Petri dishes and kept 
out of doors at a time when weather conditions were favourable to the 
development of mildew. On each successive day two slides were moistened 
with distilled water and twenty-four hours later these were examined for 
germination. The results from three experiments of this kind are given in 
Table 3, the figures being the average percentage germination of the 
conidia for each pair of slides. 


Ii6 Transactions British Mycological Society 

Table 3. Viability of mildew conidia 

% germination after exposure for 

I day 

2 days 

3 days 

4 days 

5 days 

6 days 

7 days 

8 days 

Exp. I 









Exp. II 








Exp. Ill 








Under the prevailing conditions, viability ceased after the sixth or 
seventh day. It is clear therefore that while conidia remain viable long 
enough to allow of their being widely dispersed by wind, they cannot 
transmit the disease over a lengthy time interval. 

E. Transmission by weeds 

Bremia Lactucae occurs on a large number of composite genera such as 
Sonchus^ Tragopogon^ Crepis^ Senecio and Hieracium^ and. these include many 
common species of weeds which may provide a source of infection of the 
cultivated lettuce. 

In the autumn of 1940 seeds were collected from the following composite 
Sonchus oleraceus^ Tragopogon^ Hieracium Pilosella^ H, murorum^ 
Lactuca muralis^ i. Scariola^ L, salignay Senecio Jacohaea and S. vulgaris. Seeds 
of these were sown under glass in September 1940 and grown to the three- 
or four-leaf stage, after which they were sprayed with a suspension of 
Bremia spores from cultivated lettuce by means of an atomizer. Lactuca 
saligna and L, Scariola only developed mildew lesions. A repetition of the 
experiment gave the same result, and the infections were successfully 
transferred from the wild species of Lactuca back to healthy cultivated 

Later, mildew was found occurring naturally on Senecio Jacohaea and this 
was transferred to S, vulgaris seedlings but it failed to infect the cultivated 
lettuce. Bremia found on Senecio vulgaris in the field behaved in a similar 
way, i.e. it infected S. Jacohaea but not the cultivated lettuce. 

Bremia^ therefore, though morphologically uniform, has various bio- 
logical races. These different races show some specific choice of a host, 
e.g. Lactuca muralis not infected by conidia taken from i. saligna^ 
L, Scariohy or the cultivated lettuce. 

It appears therefore that, of the wild hosts examinedy L. Scariola and 
L. saligna are the only ones which are capable of transmitting mildew to 
lettuce. As both these are uncommon in England, they cannot be of any 
importance in this connexion. 

The conclusions to be drawn from sections C, D and E are on the whole 
negative and therefore the only plausible hypothesis is that each lettuce 
crop is infected from a previous one. The implications of this will be 
discussed later. 

F. Importance of the disease 

Repeated observations of mildewed plants show that they become more 
susceptible to secondary attacks of the more important Botrytis disease. 
Mildew lesions on the outer leaves of more or less fully grown lettuce plants 

Downy Mildew Disease of Cultivated Lettuce. H. Wild 1 17 

become senescent and Botrytis invades the leaves from these areas. Usually, 
however, mildewed plants do not succumb, but mature in due course. 
Damage therefore is to be assessed by comparing the performance in 
the field of mildewed seedlings with that of uninfected seedlings of the 
same age and cultural treatment. This type of experiment would not have 
much practical significance if the grower were able to discard all infected 
seedlings when planting out, but mildew is often so widespread in the 
frame that selection is not possible, and plants with very small or incipient 
lesions could not be sorted out. 

Eight plots of lettuce were planted out from Dutch lights on 28 February 
1941, each plot consisting of four rows of fifteen plants. The odd and even 
plots consisted of mildew-free and mildewed seedlings respectively, and all 
the plants were, as far as could be seen, free from infection by Botrytis and 
Rhizoctonia^ Care was taken to see that all the plants were of uniform size. 
The survival figures for each plot were noted on 30 April and 2 June 1941, 
and these are shown in Table 4. 

Table 4, Effect of mildew on survival of lettuce 

Plants per plot 



28 Feb. 

30 Apr. 

2 June 


No mildew 










No mildew 










No mildew 










No mildew 










Free from mildew 








The presence of mildew thus causes a material increase in the mortality 
of the plants, due in this experiment as usual to Botrytis cinerea. 

The cutting record of this crop is shown in Table 5. 

Table 5, Numbers of lettuce matured at various dates 

No. of lettuce cut 




6 June 

12 June 

16 June 

19 June 



No mildew 





. 2'" 







■'2 /;■ 


No mildew 





' 2 ■ 









No mildew 









' 10 



t' ■' 


No mildew 





'2 ' 









Free from mildew 












By the method of linear interpolation, one can determine approximately 
the dates at which 75 % of the two lots of plants matured. These are 8 and 
14 June for the plants from non-mildewed and mildewed seedlings 


ii8 Transactions British Mycological Society 

respectively. This represents a check to maturing of about six days, which 
is an item of importance to the growers of spring lettuce. 

G. Control by ]vieans of resistant varieties 

Workers in several countries have claimed that lettuce shows varietal 
resistance to mildew. Jagger and Chandler (1932) demonstrated that 
immunity is a Mendelian dominant and named eight resistant varieties. 
According to Macpherson (1932) the English varieties Loos Tennis Ball, 
Rosy Spring, and May Queen are resistant. 

For the purposes of testing, seedlings three weeks old and of comparable 
size were evenly spaced in pots. There were ten seedlinp per pot and ten 
pots of each variety. They were then inoculated with mildew spores under 
identical conditions, as follows. A suspension of fresh spores was put in a 
type H 3 Aerograph spray gun, constant pressure provided by a hand pump 
supplying air at 15 Ib./ and the nozzle pointed horizontally at the 
seedlings with its tip at a distance of 9 in. from the near edge of the pot. 
Each pot was sprayed for one second. 

Table 6 gives the percentage of infected plants in an experiment with 
nine commonly grown varieties. 

Tabled. Varietal susceptibility to mildew 
Variety % infection Variety 

Lees’ Immense i*i Cheshimt Early Giant 

Trocadero 2*5 Arctic King 

Market Favourite 6*3 Blackpool 

Feltham King 12*9 Early French Frame 

Imperial 15*0 

On analysing the data obtained in this experiment by the method of 
Bliss (1937, 1938)5 the following conclusions were drawn: 

(1) Lees’ Immense is significantly more resistant than Feltham King and 
all other varieties below it in the table. 

(2) All the varieties are significantly more resistant than Early French 

(3) Trocadero and Market Favourite are significantly more resistant 
than all the remaining varieties except Feltham King and Imperial. 

In experiments which will be described later, the varieties Trocadero 
and Imperial, which would be classed respectively as resistant and 
moderately resistant on the basis of the results in Table 6, gave over 90 % 
of infected plants. Thus it is seen that, although there is a measure of 
variation in resistance between varieties, all those used by the writer have 
shown heavy infection under conditions favourable to the development of 
mildew. The choice of resistant varieties is not likely therefore to eliminate 
infection by mildew. 


That high humidity favours the disease is unquestioned and this was borne 
out by my experience in inoculating lettuce seedlings with mildew through- 
out this work. Growers accept tins view and accordingly give as much 

% infection 

Downy Mildew Disease of Cultivated Lettuce. H. Wild 119 

ventilation as the atmospheric conditions permit. Thus it is good practice, 
in the management of frames, to remove the covers completely on dry 
non-frosty days, and to allow a certain amount of ventilation at night, in 
all but severe weather, by propping up the covers. It is true that other 
considerations apply, e.g. this procedure is believed to give some protection 
against Botrytis disease. Also complete removal of lights whenever possible 
allows of better illumination, so that hardier, stockier, more frost-resistant 
seedlings are produced. 

Though so much is accepted, there is little quantitative evidence that 
humidity influences the spread of mildew; consequently the following 
experiment was devised. Two pots of similar size, each containing ten 
seedlings heavily infected with mildew were placed in the centre of neigh- 
bouring frames. Twenty pots, each containing ten healthy, three-week 
Imperial seedlings were placed round these pots at a distance of 2 ft. 
Both frames were then kept shut and one watered every day, the other not 
at all. After twenty- three days mildew lesions were noticed and the per- 
centage infection determined. That in the watered frame was 61*4 % and 
that in the unwatered frame was 1 1*3 %■. Between 27 March and 9 April 
1941 the experiment was repeated and the results were: 

Watered frame 38*6 % infection 

Unwatered frame o-o% infection 

These results are very clear-cut but though the humidity of the watered 
frame was certainly higher than that of the dry one some of the spread of 
the disease may have been due to the draughts and splashing caused by 

Whether the opening of the frames be good or bad from the point of 
view of draughts, it is certain that during spells of very humid, muggy 
weather opening of the frames cannot reduce the humidity to the safety 
level. The fact that mildew does appear in quantity both in frames and out 
of doors, especially in mild wet winters, is evidence that the ventilation 
method, even if it is useful, does not give adequate control under conditions 
which are highly favourable to the disease. 

I. Fungicidal CONTROL 

Experiments on this subject were carried out over the period November 
1939 April 1941. The general plan was as follows. For each treatment 
ten pots were set up, each containing ten three- week seedlings. These were 
hardened off and then sprayed or dusted with the appropriate fungicide. 
Next day they were inoculated with a suspension oi Bremia conidia. The 
inoculum was prepared in tap water by brushing off spores from fresh 
mildew lesions with a wet caniel-hair brush. Examination of a drop under 
the microscope showed whether it was concentrated enough, and usually it 
was filtered through butter muslin to remove conidiophores and dirt. 

In 1939-40 the inoculations and sprays were applied with a small scent- 
spray atomizer but in 1 940-1 a type HS Aerograph spray gun with a 

! 120 Transactions Bntish Mycological Society 

^ de Vilbiss nozzle was substituted since it gave a more constant and 

finely divided spray. The air pressure of 15-20 Ib./sqin. required for the 
operation of the atomizer or spray gun was provided by a small hand 
pump. Dusts were applied with an apparatus specially constructed to give 
an even flow of dust into a chamber of known area containing the pots to be 
: dusted. 

1 The pots of seedlings, after treatment with fungicide and spraying with 

I spores, were randomized in a cold frame which was kept closed. After 

some days, varying in number with the temperature, but usually about 
twelve, conidia had developed on the leaves. Each seedling, which had by 
this time at least four leaves including the cotyledons, was examined and the 
number of infected and uninfected leaves noted. At the same time the 
amount of phytocidal damage wa;s assessed for each pot according to an 
arbitrary scale, the figures o, i, 2, 3 corresponding respectively to 'no 
I damage’, 'just visible flecking’, 'fairly general spotting’ and 'development 

of considerable necrotic areas’. The highest figure obtainable in any 
experiment of ten pots was therefore thirty. Records of maximum and 
minimum temperatures were kept throughout the experiments. 

Experiment I ^ begun on 12 November 1939, contained nine treatments: 

(1) 5 % red copper oxide with talc as filler and applied as a dust at the 
rate of 0-25 oz./sq.yd. 

(2) 5 % rod copper oxide with Folosan (pentachloronitrobenzene) as 
filler, 0-25 oz./sq.yd. 

(3) I lb. red popper oxide, \ gal. cotton-seed oil, and i quart of lethalate 
spreader made up in 50 gal. of water. 

(4) 2:2: 40 Bordeaux mixture as a spray. 

(5) Folosan as a dust at 0*25 oz./sq.yd. 

(6) Folosan with lime as filler at 0-25 oz./sq.yd. 

(7) Flowers of sulphur as a dust at 0*25 oz./sq.yd. 

(8) Pure calcium hydroxide dust at 0*25 oz./sq.yd. 

(9) Control. 

Bordeaux mixture was included because of its successful use at this 
strength for the control of mildew by Osmun (1934), the copper oxide 
sprays and dusts because of the work done by Clayton ( 1 938 b) in the control 
of downy mildew of tobacco, and the Folosan because of its control of 
Botrytis disease of lettuce as demonstrated by Smieton and Brown (1940) . 
Lime was included because some growers use it in the hope of reducing 

The results of this experiment are given in Table 7 and may be sum- 
marized as follows : 

(1) Lime is ofno value. 

(2) Sulphur, though it causes no injury, gives poor protection. 

(3) Folosan, with talc filler, gives fair control and is safe; on the other 
hand, the incorporation of lime as filler more or less completely takes away 
its fungicidal value. 

(4) All the preparations containing copper give a pronounced measure 
of control, but those containing copper oxide (GugO) have rather a pro- 
nounced phytocidal effect. The best control is given by 2 : 2 : 40 Bordeaux 
•.mixture.' -'A 

Downy Mildew Disease of Cultivated Lettuce. H. Wild 121 

Table 7. Effect of fungicides on the control of lettuce mildew 




A ^ 





% infected 


% infected 


(i) Copper oxide 4* talc 






(2) Copper oxide +Folosan 






(3) Copper oxide spray 






(4) 2:2: 40 Bordeaux 





2' ; 

(5) Folosan 






(6) Folosan lime 






(7) Lime dust 






(8) Sulphur dust 






(9) Controls 






Average minimum temperature =41*0° F. 

Experiment II. This was essentially a repetition, in duplicate frames, of 
the preceding. It was begun on 19 December 1939, but was not concluded 
until 25 January 1940 on account of the very slow development of mildew. 
During this period the exceptionally cold spell of the winter of 1939-40 
began. This led to considerable winter-killing by frost, in spite of the 
protection afforded by the frames. The data of Table 8 show that this 
effect was accentuated by the use of fungicides, and in particular by certain 
of the copper-containing preparations. Of these, dilute Bordeaux mixture 
was, as in Table 7, the least damaging. 

Table 8. Mortality of lettuce seedlings due to frost 


Exp. A 
% mortality 

Exp. B 
% mortality 

(i) Copper oxide + talc 



(2) Copper oxide 4- Folosan 



(3) Copper o?dde spray 



(4) 2 : 2 : 40 Bordeaux 


• 42 • 

(5) Folosan 4“ talc 



(6) Folosan 4- lime 



(7) Lime 



(8) Sulphur 



(9) Control 



Av. min. temp* 

26-5° F. 

25-2® F* 

In further experiments during 1 940-1 sprays only were used, and the 
list of these is as follows : 

( i ) Cuprous oxide. ‘Cuprocide 54’, a commercial preparation containing 
50% CU2O with incorporated spreader, was used at strengths of 2, i, 

(2) Bordeaux mixture of compositions i : i : 40, 2 : 2 : 40 and 4 : 4 : 40. 

Excess-Lime Bordeaux (2 : 10 : 40). The fungicidal efficiency of this 
preparation has been demonstrated by Grubb (1924). 

(4) Sodium orthophosphate-copper sulphate spray, Bruno (1940) gives the 
formula for a copper fungicide as : 1 kg. copper sulphate in 50 L of water, 
and I •25-1 *30 kg. sodium orthophosphate in 20-30 1 . of water, made up to 
100 1 . in water. He claims that it fe more dispersible than Bordeaux 
mixture. This spray contains approximately the amount of copper present 
in a 4 : 4 : 40 Bordeaux mixture. Tor use on lettuce it was reduced to half 
the strength advised by Bruno to make it comparable with a 2 : 2 : 40 
Bordeaux mixture. Sodium orthophosphate, unlike lime, dissolves com- 
pletely in water, and the precipitate formed when copper sulphate is added 


Transactions British Mycological Society 

is fin ely divided and flocculent. This fungicide therefore gives a more even 
dispersion of small particles over the leaf surface than does Bordeaux mixture. 

(5) Bouisol at the rate of i lb. to 30 gal. of water. At first the spreader 
Agral 2 (0-05 %) was added to the Bouisol but was later left out as it was 
found to increase damage. 

(6) Salicylanilide at the rate of 10 0Z./40 gal. of water with 0-05 % Agral 2 
as a spreader. 

Table 9 







Date: From 

13. X. 40 

16. X. 40 


xi. 40 


xi. 40 

28. xi. 40 



I , xi. 40 


xi. 40 


, i. 41 

17. i. 41 

Average min. temp. ° F. 









KD. % in£KD. % inf! E.D, 

. %inf. 


%inf, E.D. %inf: 

i strength Cuprocide 

0 20*6 






1 strength Cuprocide 

I 13 

6 i8*i 





3*5 0 

Full strength Cuprocide 

3 21*8 

I i8*7 




/2' , ■■ 

6 0 

I : I : 40 Bordeaux mixture 

2 7*1 

0 2*1 





II 0 

2 : 2 : 40 Bordeaux mixture 

5 I 

2 I 





12 0 

4 : 4 : 40 Bordeaux mixture 

5 i’2 

3 0 





14 0 

Excess-Lime Bordeaux 

— — 




'■ 2 

“ ' — 

Orthophosphate-copper spray 

— . — 

— ^ — 

, — . : 


— ' 


— — 

— ' 



— ■ 


— •_ 

Bouisol + Agral 2 

' — — 

— ' — 





23 0 

l^alicylanilide -h Agral 2 

— — 







— ' — 


0 53 

0 46*8 




0 10 






Date: From 


28. ii. 41 


Hi. 41 


iv. 41 


25. ii. 41 

18. Hi. 41 


iv. 41 


iv. 41 

Average min. temp. ° F. 





^ .A. ■ _ 

A ^ 


E.D. % in£E.D. %inf. E.D. 

% inf. E.D. 


J strength Cuprocide 

— — 

■ ' 




1 strength Cuprocide 

— — 

' — 


Full strength Cuprocide 

2 0 

0 0 





I : I : 40 Bordeaux mixture 

. 4*5 0 

0*5 0 

I *5 

" 2 



2 : 2 : 40 Bordeaux mixture 

3*5 0 

0 2 





4 : 4 : 40 Bordeaux mixture 

4 I 

0*5 i*i 





Excess-Lime Bordeaux 

0 I 

0 2 


' ,1' 



Orthophosphate-copper spray 

0*5 I 

0 I 






3 0 

0 0 




, 12 ' 

Bouisol -j- Agral 2 

2*5 0 

Salicylanilide-j- Agral 2 


— — ^ 

— ■ 

, — ' ■ . : 






0 6i*9 





E.D. estimated damage. 

Table 9 shows the relation between fungicidal efficiency^ damage, and 
average minimum temperature. The following are the conclusions from 
this table. 

(1) The amount of damage increases as the average minimum tem- 
perature falls. 

(2) The damage varies with the fungicide. 

(3) As the strength of the Bordeaux increases there is no marked increase 
either in fungicidal effect or phytocidal damage though on the whole 
2 : 2 : 40 Bordeaux gives the best control. 

Downy Mildew Disease of Cultivated Lettuce, H. Wild 123 

(4) Cuprocide 54 somewhat safer at low temperatures than Bordeaux 
2:2: 40 and in addition it is just as effective as a fungicide. 

(5) Bouisol at the strength used (i lb. to 30 gal. of water) has no 
advantage over Bordeaux as a control and causes more damage. 

(6) 2 : 10 : 40 Excess-Lime Bordeaux causes less damage than 2 : 2 : 40 
Bordeaux mixture but its efficiency as a fungicide may be a little less. 

(7) The sodium orthophosphate-copper sulphate spray, over the short 
period during which it was used, proved as efficient as 2 : 2 : 40 Bordeaux 
and also caused less damage. 

(8) Salicylanilide + Agral 2 (5 %) gives a poor control of mildew. 

The experiments so far described were carried out under conditions 

which were highly favourable to the development of mildew. In the course 
of the work three experiments under ordinary conditions of practice, except 
that the plants were uniformly inoculated at a young stage by spraying 
with spores, gave the results which are summarized below. 

Experiment i, with fifteen Dutch lights each containing forty-eight 
Imperial seedlings (23 Feb. -3 April 1940). Treatments: Control; Cuprocide 
54 (|- lb. to 50 gal. water) ; Bordeaux mixture i : i : 40, 2 : 2 : 40 (with or 
without Folosan), 4:4: 40. Two sprayings were applied. All treatments 
gave good control of mildew, the controls showing considerable infection 
in the young stages. The experiment was concluded when the plants were 
about half-grown, at which time there was no evidence of difference 
between controls and treatment in respect of numbers of survivors or 
weight of plants. This was apart from some check to growth where Folosan 
had been used. 

Experiment 2, with six Dutch lights (4 Nov. 1940-13 May 1941). Treat- 
ments : Control; Bordeaux mixture 2 : 2 : 40 and 4:4: 40. Three sprayings 
were applied. Conditions in early winter were not favourable to severe 
infestation of the controls. At the time of maturity there was no difference 
in any respect between treated and untreated plants. 

Experiment 3, with sixteen Dutch lights (17 Dec. 1940-12 June 1941). 
Treatments: Control; Bordeaux 2 : 2 : 40; Cuprocide 54 (i lb. to 50 gal. 
water); Bouisol (i lb. to 30 gal. water). Two sprayings were applied. By 
mid-February heavy infestation of the untreated frames was shown, the 
treatments and especially 2:2: 40 Bordeaux showing good control. 
Seventy-two plants from each treatment were set out in replicated plots in 
the open ground on 18 Feb. and carried to maturity. The differences in 
survival and in average mature weight were negligible, but there was a 
delay of five days in the maturing of the untreated plants. This amount 
may be of significance to the practical grower, at least at certain seasons. 

Attempts to use the vapour method which has proved successful in the 
control of tobacco mildew (blue mould), viz. to use paradichlorbenzene as 
recommended by Clayton (i938<2, i) or benzene mixed with lubricating 
oil as recommended by Wolf et aL (1939, 1940) have given very unsatis- 
factory control of lettuce mildew and do not appear at all promising. 

J. Effect of pH on germination of Bremia GomDix 
The preliminary experiments in the section on fungicidal control include 
hydrated lime, as lettuce growers consider that it gives some control of 

124 Transactions British Mycological Society 

lettuce mildew. The results obtained show that lime is of no value whatever 
as a fungicide in this instance, and infection on lime-dusted plants is at 
least as heavy as on controls, if not heavier, the comparative percentage 
infections being: ‘ 

Lime, 927 %, 98 %. Control, 87-9 %, 93 %. 

Laboratory experiments on the germination of Bremia spores on lime- 
dusted slides showed that, as expected, lime dust had no effect on the 
germination. Both controls and dusted slides had germinations varying 
between seventy and eighty per cent. This shows that Bremia conidia are 
able to germinate in alkahne solutions. The effect of pH. on germination is 
shown more fully in Table 10. 

Table 10. Effect of pH on the germination of conidia 

% germination 


Exp. A 

Exp. B 
















Both experiments agree in showing that germination is reduced in 
distinctly acid solutions but takes place freely in alkaline solutions. The pH 
of saturated lime solution however is greater than 9*6 and so is outside their 
range. Titration with hydrochloric acid by the standard method gave the 
normality of saturated lime solution as .A 734 - Further experiments showed 
that germination falls off sharply beyond this point and that it is inhibited 
in jVy 10 sodium hydroxide. On the acid side germination begins to be 
affected in concentrations of HCl exceeding i in 10,000 jV. 

K. Summary 

1 . Lettuce seedlings, grown from both ordinary commercial seed and 
seed from mother plants known to be infected with mildew, almost 
invariably failed to show symptoms of the disease under conditions 

. favourable to mildew development. There was thus negligible evidence of 
the seed-borne transmission of mildew. 

2. Oospores were not found by the microscopical examination of debris 
from infected plants, and seedlings grown in soil containing a high pro- 
portion of such debris for a period considered to be adequate, under 
favourable conditions for the development of mildew, did not become 
infected. The transmission of the disease by contaminated soil is therefore 

3. Attempts to transmit mildew from composite weeds to lettuce were 
successful only with some wild species of Lactuca. These species are probably 
too rare to be important in the spread of the disease. 

4. The survival of mildewed lettuce was lower, and their date of 
maturity later, than that of healthy plants. This, together with the dis- 
figurement caused by the disease, gives it its commercial importance. 

Downy Mildew Disease of Cultivated Lettuce. H. Wild 125 

5. A number of commercial lettuce varieties commonly grown in 
England showed varietal differences in their susceptibility to the pathogen 
but these differences were not sufficiently marked to make the exclusive 
cultivation of the more resistant varieties worth while as a control measure. 

6. 2 : 2 : 40 Bordeaux mixture, Excess-Lime Bordeaux (2 : 10 : 40), an 
orthophosphate copper spray, and Cuprocide 54 (i lb. to 50 gal.) all gave 
adequate protection from mildew disease but sprays containing no copper 
such as lime sulphur and pentachloronitrobenzene did not. All the copper- 
containing sprays are liable to produce phytocidal damage, particularly in 
cold weather. 

7. Fungicidal vapours such as benzol and paradichlorbenzene were 
troublesome to use and gave a poor control of lettuce mildew. 

8. Brenda conidia under favourable temperature conditions germinated 
readily in saturated lime solution and were also found to germinate more 
readily in slightly alkaline than in slightly acid solution. 

I wish to thank Prof W. Brown, F.R.S., who suggested this investigation 
and under whose supervision it was carried out. 


Baudys, E. (1935). Lettuce or salad mould. Leaji. phytopath. Sect. Reg. Agric. Exp. Sta.^ 
Brnoy XGiir, 2 pp. 

Bliss, G. (1937). The analysis of field experimental data expressed in percentages. Plant 
Protection, xii, 67-77. 

Bliss, C. (1938). The transformation of percentages for use in the analysis of variance. 
Ohio J. Sci. XXXVIII, 9-12. 

Bruno, A. (1940). A new formula for a copper mixture. C.R. Acad. Agric. Fr. xxvi, 

454 ” 7 * 

Clayton, E. E. (1938^2). Paradichlorbenzene as a control of blue mould of tobacco. 
Science, N.S. lxxxviii, 2272, p. 56. 

Clayton, E. E. etal. (19386). Control of the blue mould (downy mildew) of tobacco by 
spraying. Tech. Bull. U.S. Dep. Agric. 650, 23 pp. 

DE Bary, a. (1863). Recherches sur le , developpement de quelques champignons 
parasites. Ann. Sci. Nat. Bot. (Ser. 4), xx, 5-148. 

Grubb, N. H. (1924). Tests of fungicides on apple trees. J. Pomol. Hort. Sci. in, 157-62. 
Jagger, I. C. & Chandler, N. (1932). Physiologic forms of Bremia Lactucae RegeL 
Phytopath. 18-19. 

Lavrov, N. N. (1932). Key to plant parasites of cultivated plants of Siberia. Pt. i. 
Abs. in Rev. Appl. Mycol. (1933), xii, 306-7. 

Magpherson, N, j. (1932). The cultivation of lettuce under glass with special reference to 
varietal resistance to downy mildew, J. Minist. Agric. xxxviii, 998-1003. 

OsMUN, A. V. (1934). Rep. Dep. Bot. Mass. Agric. Exp. Sta. pp. 23-7. 

Sghweizer, j. (1919). Die kleinen Arten bei Bremia Lactucae RegeL Verh. Thuring. 
naturf. Ges. icsm, 

Smieton, M. j. & Brown, W. Botrytis disease of lettuce, its relation to damping- 

oflf and mildew, and its control by pentachloronitrobenzene dust. Jnw. appl. Biol. 
xxvii, 489-501. 

Smith, W. G. (1884). Resting spores of Peromspora gangliformis Berk. Gard. Chron. xxi, 

WoLF, F. A. et al. (1939). Field studies on the concentration of benzol vapour as used to 
control downy mildew of tohdicco. Phytopath, 'xxix, 177-87. 

’Wo'LF,P. A. et al. (1940). Volatile fungicides. Phytopath, 'kxk, 21^-2 J. 

{Accepted for publication 28 December 

[ 126 ] 



Technical Director^ Noble Mushrooms Ltd.^ Yaxley^ Peterborough 

A new species of Verticillium attacking the brown strain of the cultivated 
mushroom, Psalliota (Agaricus) campestris (L.) Fr., was recently described in 
Copenhagen by Treschow (1941), who named it Verticillium Psalliotae. The 
same species was noticed at Yaxley in January 1946, attacking the white 
strain of the cultivated mushroom; and other specimens have since been 
seen at Thornham, Norfolk. On 29 January 1946, during the course of 
routine examinations of diseased mushrooms which modern commercial 
practice demands (Atkins, 1945), one specimen with a light brown, roughly 
circular patch on the pileus was found, as expected, to be attacked by a 
Verticillium, but the species differed radically from F. Ware 

(1933) in that the conidia were invariably pointed at each end, and they 
were borne transversally on the phialides. Commercial practice com- 
pelled the removal of the diseased mushroom from the bed, but cultures 
were made in the laboratory. Three more similarly infected mushrooms 
were found in succeeding weeks, and detailed examination by Mr McG. 
Bulloch and myself revealed close similarity to V: Psalliotae, which Treschow 
found in the summer of 1939. 

Treschow (1941) stated that the pathogenicity of both F. Malthousei and 
F. Psalliotae was confined to the brown strain. This contention is certainly 
invalid with regard to F Malthousei, as Ware (1933) has demonstrated; 
and from the appearances at Yaxley it appears that the white strain is also 
attacked by F. Psalliotae. We sent cultures of our fungus to Dr Treschow, 
and after a preliminary examination he tells me he is unable to find any 
difference between it and F. Psalliotae. He found that the appearance of 
the strain, the position of the branches, and the size, as well as the character- 
istic red colouring matter formed in the substratum, were identical. 

White mushrooms at Yaxley have been successfully inoculated with the 
disease by applying drops from a washing in sterile water of a malt agar 
slope culture. Vigorous growth of mycelium was visible to the naked eye 
after forty-eight hours at 20"^ C. 

The disease is now being investigated further by Dr G. J. LaTouche, 
Microbiologist with the Mushroom Research Association at Yaxley. 
Preliminary observations reveal that the mould, when grown on 2 % malt 
agar {pH. 5*5) at fluctuating laboratory temperatures, produced an 
abundant floccose snow-white aerial mycelium which was compacted to 
form a thin skin at the surface of the medium. It developed a pigment 
(Van Dyke Red, Ridgway) in the medium, as Treschow found. Aerial 

N ttticiViivm Fred. C. Atkins 127 

mycelium from a three-weeks-old culture on malt agar, mounted in lacto- 
phenol and cotton blue, revealed the following characteristics : 

Hyphae septate, branched, about i p. broad. Fertile hyphae bearing 
more than four verticils of phialides, which in turn vary in number from 
two to five, measure i2*5~3i-5 ju- long by a maximum of i /x broad, and 
taper to a fine point. Conidia aseptate, more or less straight on the 
attached side and somewhat convex on the other, pointed at both ends, 
4*3-i2-97xx I-75--2-I5/X broad (compared with Treschow’s measurements 
of 6-10*5 /XX 2--3-5^). 


Atkins, F. G. (1945). on mushrooms. Mushroom Growers^ Association^ Midlands 

Group Publication^ Taxley, Peterborough, pp. 9-25. 

Treschow, G. (1941). The Verticillium diseases of cultivated mushrooms. Dansk Bot. Ark. 
XI, 1-31. 

Ware, W. M. (1933). A disease of cultivated mushrooms caused by Verticillium 
Malthousei sp . nov . .dw/z. Zoni., xlvii, 763-85. 

[Accepted for publication 31 December 1946) 

[ 128 ] 




The Freshwater Biological Laboratory, Wray Castle, Ambleside, and 
Department of Botany, Birkbeck College, London 

(With Plate XI and 4 Text-figures) 

The genus ZyS^'^^^zidium was erected by Lowenthal (1905) for an epibiotic 
operculate clxytrid, distinguished from Chytridium by possessing a resting 
spore formed as the result of a sexual process. The type of sexual reproduc- 
tion exhibited by is well known. A small male thallus puts 

out a conjugation tube which grows until it makes lateral contact with the 
wall of a larger female thallus. Fusion occurs and the contents of the male 
pass into the female, which later becomes the resting spore. In addition to 
LowenthaFs observations on Z- it has been described by Scherffel 

(1925) and Domjan (1936). verrucosum Geitler (1942) has an identical 
sexual process (Text-fig. if). 

This type of sexuaf reproduction has again been observed in Chytridium 
Characii Scherffel (1925). The mature resting spore is ovoid, with its long 
axis at right angles to the host cell. The outer layer of the wall is thicker at 
the base and apex than in the middle region, and in surface view is covered 
by longitudinal rows of elongate warts (Text-fig. T ^). Although the 
method of formation of this resting spore is so strikingly like that of 
Zygorhizidium, the exact position of Chytridium Characii remains obscure 
since no zoosporangia were observed. The conjugation tube in the three 
organisms mentioned above is characterized by remaining equally 
cylindrical throughout its length. Scherffel (1925) describes Chytridiuml 
Spirotaenia, in which the conjugation tube swells at its point of contact with 
the female, and becomes club-shaped (Text-fig. ib). Thus when the male 
and female thalli are close together, the narrow middle portion is lost, and 
the conjugation tube forms an irregular vesicular structure (Text-fig. i^). 
Once again the exact affinities of this organism are unknown, since 
dehiscence of the zoosporangium was never seen. The structure of the 
resting spore wall, with its outer surface covered with spines, consisting of 
refractive wall material, somewhat resembles Zyg^'^^izidium verrucosum, and 
the type of sexuality may be regarded as a variant of that found in Zyi^” 

A few examples of >^. Willei, hitherto unknown from this country, were 
found growing on Mougeotia sp. in a collection from Montreal Park Lake, 
Sevenoaks, Kent, England, in March 1945. Little doubt remains as to the 
identity of this fungus a^^^ dehiscence of the zoosporangium was not 

130 Transactions British My cological Society 

seen, since the shape of the sporangia and the peculiar method of sexual 
reproduction agree perfectly with Lowenthal’s original description of 
Zygorhizidium Willei on Cylindrocystis Brebissonii, from Norway. 

The fungus is parasitic, but rarely produces more than a slight dis- 
organization of the host chloroplast. Large sporangia 17/x in diameter 
may bring about a curvature of the Mougeotia cell as reported previously by 
ScherflPel (1925), but this is not a constant feature, and dwarf sporangia 
(5”6*6/^ in diameter) caused no distortion of the filament. The intra- 
matrical rhizoidal system consists of a knob-like structure apparently 
devoid of rhizoids (Text-fig. 2b). In some specimens it seemed to be absent, 
but more probably it was merely obscured by the chloroplast of the host 
cell. Zoospores were observed only once, and were of the usual chytridi- 
aceous type. 

After the material had been left in the laboratory for about four weeks, 
sexually formed resting spores appeared (Text-fig. 2f~h). The male thallus 
(PI. XI, fig. 2), is ovoid and possesses an endobiotic knob devoid of 
rhizoids. The conjugation tube from the male thallus often branches, and 
may reach a length of 33 /x. Contact is made with a larger subspherical 
receptive thallus whose endobiotic knob also lacks rhizoids. In several 
instances two conjugation tubes approached the receptive thallus (Text-fig. 
2 A), but the one which had fused with it could be detected by its lack of 
contents. The resting spore is spherical, 1 1 in diameter, with a thick 
smooth wall, brownish in colour, surrounding the central highly refractive 
globular contents; germination was not observed. 

In late August 1946 a chytrid, at first believed to be Chytridium? Spiro- 
taenia Scherffel, was found parasitizing Spirotaenia condensata Breb. in the 
Clay Pond, Wray Castle. The fungus appears to be specific to its host, 
since other Conjugales, Mougeotia^ Zyg'^^'^^ Closterium although 
present, were not attacked. The epibiotic sporangium is usually broadly 
ovoid, very rarely spherical, with its longer axis paralfel to the algal wall 
(Text-fig. ^b~f). A small conical protuberance, probably representing an 
unexpanded portion of the original zoospore case, can usually be found on 
the upper surface of each sporangium (Text-fig. 3 c). The contents of the 
latter are in young stages mainly centralized, with a few scattered globules 
in the hyaline peripheral region. Later the content becomes evenly 
granular, and the subsequent changes in the protoplasm leading to the 
formation of zoospores are similar to those described for the majority of 
chytrids. The rhizoidal system where visible is not extensive and consists of 
a tuft of branched structures arising close together, so that no distinct main 
rhizoidal axis is produced (Text-fig. 3^, d). The mature sporangia vary 
from 25 to 63 /X broad X 16 to 27ft high; a few extremely small ones rather 
more spherical were encountered 8-“i5 fc broad x 9-13 pc high. The number 
of zoospores produced in a mature sporangium varies according to its size. 
A small one liberates about fifteen zoospores, whereas up to one hundred 
are formed in a large sporangium. The sporangium wall gradually 
deliquesces, and two oppositely placed, broad dehiscence pores appear 
(T ext-fig. 3 f) . The uniflagellate zoospores emerge singly, and swim 
away with a smooth gliding movement. They are spherical, 2*6pc in 


Transactions British Mycologicat Society 

diameter, with a conspicuous anterior oil globule and a darker area to one 
side of it. 

The resting spore is produced after a sexual process identical with that 
found in The spherical male thallus puts out a narrow 

conjugation tube about 2 fi in diameter, up to 38/4 long, which grows until 
it reaches a female thallus (Text-figs. 3^, 4A-A:, n and PL XI, fig. 5). The 
latter is also spherical, slightly larger than the male and having a broader 
base. The rhizoids of these thalli can rarely be distinguished. Presumably, 
following fusion, the wall of the female thickens (up to 3^6 diameter) and 
columnar bands of highly refractive wall material develop. These extra 
thickening bands are internal to the spore wall, which retains its smooth 
outline (Text-figs. 3^, 4 A, n and PL XI, fig. 4). The mature resting spore 
is spherical, io-20/r in diameter, with granular contents; its germination 
was not observed. 

As mentioned earlier (p. 130) this chytrid shows a striking resemblance 
to Chytridium Spirotaenia, The shape of the zoosporangium, with its conical 
protuberance on the upper surface, is similar, as is also the method of 
sexual reproduction by conjugation. However, there are certain differences. 
The sporangium is much smaller, opening by a single apical or lateral 
pore, and not by two lateral oppositely placed pores as is characteristic for 
Rhizophidium columnaris. Also in Chytridium Spirotaenia the male conjugation 
tube swells at its point of contact with the female, and the outer layer of the 
resting spore wall is densely covered with broad, solid, refractive, ray-like 
protuberances, giving it an irregular outline (Text-fig. ib^c). In view of 
these differences it is suggested that these two organisms cannot be 
recognized as identical, and a new species is erected, Rhizophidium columnaris^ 
taking its name from the columnar bands of thickening on the resting 
spore wall. 

In R, columnaris^ as in Zygorhizidium^ certain aspects of its sexuality 
remain puzzling. Both Scherffel and Lowenthal describe dwarf thalli of 
Z- Willei, upon which a conjugation tube had formed, functioning as 
zoosporangia. In Rhizophidium columnaris one such specimen was observed 
(Text-fig. 4772). However, although the oil globules of the zoospores were 
delimited their actual liberation was not seen. This possibly gives further 
support to the suggestion, that the subsequent nature of the thallus 
produced from the zoospores in Zyg^'^bi^idium is determined by environ- 
mental conditions, and not due to inherent differences produced in the 
swarmers on germination of the resting spore. . 

Rhizophidium columnaris n.sp. 

Thallus monocentric, epibiotic. Sporangium broadly ovoid, with its 
longer axis parallel to the. host wall; wall smooth, colourless with a conical 
protuberance on its upper surface; large sporangia; 25-63 p, broad x 16- 
27/x high; dwarf sporangia 8-15/^ broad x 9-13 /x high; dehiscing by two 
broad, lateral, oppositely placed pores, very rarely one apical pore. 
Zoospores spherical, 2*6 ju. in diameter, uniflagellate, with a conspicuous 
anterior oil globule and a darker area laterally ; emerging singly, movement 



Text-fig. 3. Rhizophidium columnaris n.sp. a, encysted zoospores, b, young zoosporangium with 
centralized highly refractive contents; rhizoids not visible, c, young zoosporangium with 
well-developed rhizoids, the conical protuberance is visible on the upper surface, (f, mature 
sporangium, oil globules of the zoospores delimited, e, dehisced sporangium with zoospores. 
/, two empty sporangia and a mature dwarf sporangium, g, various stages in development of 
sporangia and resting spores. (All X 660, except x 1400.) 


Studies on British Chytrids. Hilda M. Canter 135 

even gliding. Rhizoidal system branched arising from an indistinct main 
axis. Resting spore sexually formed, epibiotic, spherical, 10-20/^,, wall up 
to 3 fc thick, colourless, smooth, with columnar bands of refractive material; 
central contents granular, germination not observed. Male thallus epi- 
biotic, spherical, connected to the female by a narrow cylindrical conjuga- 
tion tube 2 jC6 diameter and up to 38//, long. 

Parasitic on Spirotaenia condensata Breb., Clay Pond, Wray Castle, 

\ Rhizophidium columnaris 

Thallus monocentricus, epibioticus. Sporangium late ovoideum, 
decumbens, hyalinum, laeve, episporio una extremitate papillato; 
sporangia majora 25-63 x 16-27 H'y sporangia nana 8-1 5 x 9-13 /-t, a duobus 
poris lateralibus oppositis vel rare a singulo poro apicale dehiscentia. 
Zoosporae sphaericae, 2*6ft diam., uniflagellatae, guttula anteriore 
distincta parteque obscuriore laterale praeditae, singulatim emergentes. 
Rhizoidea ramosa ex axe parum claro oriunda. Sporae perdurantes 
epibioticae, sphaericae, 10-20 jx diam., intus granulosae, episporio ad 3 /i. 
crasso, hyaline, laeve, interne lineis radiantibus refringentibus praedito; 
germinatio non visa. Thallus masculinus epibioticus, sphaericus, tubulo 
anguste 2ja diam. et ad 38//- longo cum femine conjunctus. 

Hab. in Spirotaenia condensata Breb. parasiticus. Clay Pond, Wray Castle, 

My thanks are due to the Director of the Freshwater Biological Associa- 
tion for the use of a laboratory in which this work was done, to Miss E. M. 
Wakefield for the Latin diagnosis and especially to Prof. C. T. Ingold for 
his helpful criticism. 


Domjan, a. (1936). ‘ Vizigombas’-Adatok Szeged es Tihany Videkerol (‘ Wasserpilz’- 
daten aus der Umgebung von Szeged und Tihany). Folia cryptogam, ii (i), 8-59. 
Geitler, L. (1942). Eine neue Ghytridiale, Zygorhizidium verrucosum^ n.sp., und ihre 
Wirkung auf die Wirtszellen. Arch. Protisterik . -kcvi {i)^ 

Lowenthal, W. (1905). Weitere Untersuchungen an Ghytridiaceen. Arch. Protistenk. v, 

Sgherffel, a. (1925). Zur Sexualitat der Ghytridineen (Der ‘Beitr%e zur Kenntnis der 
Ghytridineen’, Teil I). Liii, 1-58. 

Sparrow, F. K. ( 1943) - Aquatic Phycomycetes. Ann Arbor, U.S. A. : University of Michigan 
'Press. ^ 

Zyiorkizidium Willei howtixthal a.nd Rhizophidium columnaris n.sp. 

Fig. I. Empty sporangium of ZyS^^hizidium Willei with a broad lateral dehiscence pore. A slight 
curvature of the Mougeotia cell brought about by the fungus is visible, x 1030. 

Fig. 2. Zygorhizidium Willei, male thallus with its conjugation tube, x 960. 

Fig. 3. Part of a Spirotaenia cell showing zoosporangia and resting spores of Rhizophidium colum^ 
naris. x 820. 

Fig. 4. A portion of fig. 3 more highly magnified showing two resting spores and a young 
zoosporangium. The columnar bands of thickening material are visible in the wall of the 
upper resting spore, x 1287. 

Fig. 5. PdiXt otn Spirotaenia cell with male and female thalli of Rhizophidium columnaris. {a) male 
thalli, (i>) conjugation tube, (c) female thalli. X 1 200. 

[ 136 ] 



By GEORGE SMITH, London School of Hygiene and Tropical Medicine 

Van Luijk (1938), in a paper dealing with antagonism between soil fungi, 
showed that, when Penicillium expanSum (Link.) Thom, is grown on Knop’s 
solution containing 4% sucrose, the culture filtrate has very marked 
powers of inhibiting the growth, in vitro ^ of Pythiurn de Baryanurn. In addition, 
good results were obtained in suppressing the attack of the Pythiurn on 
lucerne seedlings by treating infected soil with the active filtrate. The 
substance responsible for the inhibitory effect was not, however, isolated 
and characterized. 

Anslow, Raistrick and Smith (1943) found that patulin, previously 
isolated as a metabolic product of Penicillium patulum Bainier, is produced 
also, though in smaller yields, by various strains of P. expansum isolated 
from rotting fruits. It was demonstrated that patulin completely inhibits 
the growth of a number of species of Pythiurn at a concentration of i : 400,000, 
and that the fungistatic activity of culture filtrates from Penicillium 
expansum or P, patulum is in good agreement with their content of patulin, 
rendering it highly probable that van Luijk’s active principle was indeed 

It is now shown that patulin inhibits the growth of a number of other 
parasitic fungi, albeit at higher concentrations than that required to 
suppress the growth of species of Pythiurn, Rhizoctonia Solani Kiihn is 
completely inhibited by patulin at a concentration of i : 10,000, Sclerotium 
Rolf sit Saccardo at i : 20,000, Verticillium alb o-atrum Reinke & Berth, at 
I : lOfioofHelminthosporium monoceras Drechsler at i : 20,000, H, gramineum 
Rabenhorst at i : 20,000, AT, sativum Pamm., King & Bakke at r : 5000, 
Ceratostomella Pini Miinch. at i : '20,000, whilst Botrytis cinerea Pers. ex Fries 
is more resistant, being only partially inhibited at a concentration of 
I : 5000. It was also considered of interest to test the effect of patulin on 
Trichoderma viride Pers. ex Fries, a mould which itself produces anti-fungal 
substances (Weindling, 1932; Brian & McGowan, 1945). This species, 
like Botrytis cinerea^ is only partially inhibited at a concentration of i : 5000. 
At lower concentrations the effect of patulin is stimulatory, giving more 
abundant fructification and enhanced production of pigment. 


Cultures. Typical strains oi Rhizoctonia Solani (L.S.H.T.M. Catalogue no. 
BB192) mid Sclerotium Rolfsii (BB191) were obtained from the National 
Collection of Type Cultures. Two strains of Verticillium albo-atrum were 
received from the East Mailing Research Station, where they had been 

; Anti-fungal Substances from Moulds, George Smith 137 

h isolated from diseased hops, one (X23) from a “^progressive’ outbreak of 

j the disease, the other (X24) from a typical ^fluctuating’ infection. The 

j culture of Helminthosporium monoceras (Agiiy) was obtained in 1932 from 
C.B.S., Baarn. This species of the genus was included because it grows well, 

J and regularly produces abundant , conidia, on a variety of culture media. 

The strain of H, gramineum (Agio6) was obtained from Prof. F. T. Brooks 
in 1925 and does not produce conidia in culture. H. sativum (Ag 105) was 
also obtained from Prof. Brooks, in 1932. Cultures on a variety of media 
produce conidia only tardily and sparingly. The strain of Botrytis cinerea 
(BB9A) was isolated in this laboratory from a wilted Calceolaria seedling. 
The culture of Trichoderma viride was isolated in 1927 from cotton yarn. It 
is a fast-growing strain and produces abundant yellow pigment on potato 
dextrose agar or wort-agar. Ceratostomella Pini was obtained from Dr 
W. P* K. Findlay. 

Method of test 

The method described by van Luijk (1938), and used in this laboratory 
for the experiments with species of Pythium, is not suitable for testing 
organisms which grow only with difficulty when inoculated at the bottom 
of a tube of liquid medium. The method adopted was to make solutions of 
patulin in melted agar culture medium, pour into Petri dishes, and to sow 
each dish at a single central point with the fungus to be tested. In every 
series two or three dishes, containing the agar medium without patulin, 
were used as controls. The period of incubation varied with the fungus 
under test, but it was always at least as long as the time required for the 
control plates to be completely covered with typical growth. 

As a preliminary, the action of patulin on Pythium ultimum -w?is tested in 
this way, in order to find whether the two methods give comparable 

The agar solutions of patulin were a series of twofold dilutions from 
I ; 80,000 to I : 1,280,000. A number of solutions were made of patulin in 
Czapek-Dox solution, the concentrations being fifteen times the final 
concentrations required. These solutions were sterilized by steaming for 
fifteen minutes. Each final dilution was made by adding i ml. of the 
appropriate solution, with aseptic precautions, to a tube containing 14 ml. 
of melted Czapek agar at 45° C, The contents of the tube were well mixed 
and poured into a Petri dish. Dilutions were made in triplicate, each dish 
being inoculated with a fragment of mycelium from a young, vigorously 
growing culture of Pythium ultimum. The dishes were incubated at 24° C. 
for six days. Growth was completely inhibited at i : 320,000 and partially 
inhibited at i : 640,000. The activity of patulin, as shown by this method of 
test, is therefore of the same order as that shown by the earlier method, 
using liquid culture media. 


Rhizoctoma Solani ywm tested in the same way as Pythium, but using 
a dilution series starting at i : 5000. The results were somewhat erratic. In 
one series there was complete suppression of growth on all plates at 

j^g TvciTiscictions Btitish Mycologiccd Society 

I • 10 000 two out of three plates at both i : 20,000 and i : 40,000 showed 
no growth after thirteen days’ incubation, and there was definite retarda- 
tion of growth at i : 80,000 and i ; 160,000. In a second series there was 
again complete inhibition at i : 10,000, and on one plate at i : 20,000, but 
with little evidence of partial inhibition at higher dilutions. 

Sderotium Rolfsii. A series of tests using Czapek agar as the basal medium 
gave irregular results, but growth on the control plates, containing no 
patulin, was also erratic. The fungus grows much better on potato dextrose 
agar, and this medium was selected for a second series of tests. It was found, 
however, that solutions of patulin in potato extract turn brownish on 
heating. The pH. of the extract is approximately 6, at which reaction 
patulin is not heat-stable. In order to avoid destruction of the substance 
during sterilization, the primary dilutions were made in potato extract 
containing dextrose, previously adjusted to pH 4-5 by addition of hydro- 
chloric acid. In making the final dilutions in potato dextrose agar, the pH 
of the latter was not adjusted, since the temperature did not exceed 45° G. 
and, after addition of the patulin solution, the agar was maintained at this 
temperature for only a few seconds before being poured into a cold Petri 
dish. The dilution series used was i ; 5000 to i : 80,000 in twofold steps. 
Plates were sown with fragments of mycelium from a young culture of 
S. Rolfsii, since the sclerotia are germinated only -with difficulty in the 

There was complete inhibition at i : 20,000, with no sign of partial 
inhibition at higher dilutions. This clean fine between complete inhibition 
at I : 20,000 and normal growth, with production of abundant sclerotia, at 
I : 40,000, is in sharp contrast to the reaction of Verticillium albo-atrum. 

Verticillium albo-atrum was tested on Czapek agar, the dilution series being 
I : 5000, 1 : 10,000 and then twofold dilutions from i : 25,000 to i : 200,000. 
Both strains, X 23 and X24, gave similar results, and with each there was 
very little variation between the three plates used for each dilution. 
Inhibition was complete at i : 10,000 and there was progressively more 
abundant growth from i : 25,000 to i : 200,000. Even at the highest 
dilution, however, there was appreciable retardation of growth, as com- 
pared with the controls. 

Helminthosporium monoceras. Dilutions were made in Czapek agar, the 
series being i : 5000 to i : 160,000 in twofold steps. There was complete 
inhibition of growth at r : 10,000, and no visible growth at i : 20,000 after 
sixteen days’ incubation. At the latter dilution microscopic examination 
of the plates after three days showed that a high percentage of the conidia 
used as inoculum had germinated. Further incubation, however, resulted 
in no further growth, the germ-tubes remaining short, unbranched, and 
abnormal in appearance. There was definite retardation of growth up to 
I : 80,000. 

Helminthosporium gramineum. Potato dextrose agar was used for the tests, 
the primary dilutions being made with potato extract adjusted to pH 4-5, as 
iov Sderotium Rolfsii. The fungus was completely inhibited at a concentration 
of I : 20,000 and growth was retarded even at the highest dilution used, 

I : 160,000. In the early stages of growth all colonies showed the typical 

Anti-fungal Substances from Moulds, George Smith 139 

purplish pigmentation, but after about ten days the plates at dilutions 
I : 4O5OOO and i : 80,000 showed only small central areas of pigment, 
whilst in the controls and the r : 160,000 plates pigmentation was general. 

Helminthosporium sativum. This was tested on potato agar, at dilutions 
from I : 5000 to i : 160,000. Growth was completely inhibited at i : 5000 
and there was partial inhibition at all higher dilutions. The colonies on 
different concentrations of patulin showed very little variation as regards 
texture and tendency to sporulation. 

Ceratostomella Pini. This species also was tested on potato agar, as this has 
been found to be most favourable for production of perithecia. There was 
complete inhibition at i : 20,000 and retardation of growth up to i : 80,000. 
At 1:40,000 there was no growth after three days’ incubation, although 
in this period the colonies on the control plates had reached 3*5 cm. in 
diameter, and even after seventeen days colonies remained thin and pale 
coloured, without any suggestion of perithecia. 

Botrytis cinerea was tested in exactly the same way as Verticillium albo- 
atrum. At a concentration of patulin of i : 10,000 there was no apparent 
effect on the growth of the fungus, and at i : 5000 there was only slight 
retardation of growth. 

Trichoderma viride. Czapek agar was used for dilutions from i : 5000 to 
I : 160,000. The control plates and those containing dilutions i : 10,000 
upwards were completely covered by the mould in six days. At this stage 
the I : 5000 plates had colonies about 5 cm. in diameter, but after nine 
days even these plates were completely covered. The agar in the controls 
and the i : 5000 dilution plates remained colourless for several days and 
showed only a faint yellow colour after nine days’ incubation. At all other 
dilutions the agar was definitely pigmented after three days and rapidly 
turned deep yellow. There is no doubt that, on Czapek agar, patulin in low 
concentrations stimulates pigment production. 


AnsloWj W. K., Raistrick, H. & Smith, G. (1943). Anti-fungal substances from moulds. 
Part 1 . Patulin (anhydro-3-hydroxymethylene-tetrahydro-i :4-pyrone-2 -carboxylic 
acid), a metabolic product of Penicillium patulum Bainier and Penicillium expansum 
(Link) Thom. J. Soc, Chem, Ind.^ Lond.^'Lym, 

Brian, P. W. & McGowan, J. C. (1945). Viridin: a highly fungistatic substance pro- 
duced by CLVi, 144-5. 

Van Luijk, A. (1938). Antagonism between various micro-organisms and different 
species of the genus parasitizing upon grasses and lucerne. Med. Phytopath, 

Lab. ^ Willie Commelin Scholten^ y 'KiY, 

Weindling, R. (1932). Trichoderma lignorum as a parasite of other soil fungi. Phytopath, 
XXII, 837-45. 

[Accepted for publication 20 January 1947) 

[ 140 ] 


The method of measuring Potato Blight on the foliage proposed by a Sub- 
Committee of the Society’s Plant Pathology Committee (Moore, 1943, 
p. 34) has now been tested over several seasons by a number of observers, 
and Its practical usefulness has been satisfactorily established. Briefly, the 
method is to assess the percentage of leaf area destroyed by Blight by eye 
judgement based on the following Key, which is now presented in its 
finally revised form. 


Percentage (d.m.) 



Not seen on field 


Only a few plants affected here and there; up to one 
or two spots in 112 yd. radius 


Up to ten spots per plant, or general light spotting 


About fifty spots per plant or up to one leaflet in ten 


Nearly every leaflet with lesions, plants still retaining 
normal form: field may smell of blight, but looks ■ 
green although every plant affected 


Every plant affected and about one«half of leaf area 
destroyed by blight: field looks green flecked with 


About three-quarters of leaf area destroyed by blight: 
field looks neither predominantly brown nor green. 

In some varieties the youngest leaves escape infection 
so that green is more conspicuous than in varieties 
like King Edward, which commonly shows severe 
shoot infection 


Only a few leaves left green, but stems green 


Ail leaves dead, stems dead or dying 

In the earlier stages of a blight epidemic parts of the field sometimes 
show more advanced decay than the rest and this is often associated with 
the primary foci of the disease. Records may then be made as, say i + pf 25, 
where pf 25 means 25% in the area of the primary foci. 

Previous methods of blight measurement have adopted arbitrary scales 
of blight amounts. The present method has the advantage, that the figure 
given in each case, is a measure within fairly accurate limits of the actual 
percentage of leaf area destroyed. The error at the bottom of the scale will 
probably be less in magnitude than in the region of 50 and 75 %, but great 
accuracy is not really essential in this type of work, particularly when 
several observations are made at different dates through the season. 

The Measurement of Potato Blight 141 

It should be emphasized that the method of Potato Blight measurement 
here outlined applies only to the disease on the foliage. For disease in the 
tubers a quite different method should be adopted. For most purposes the 
percentage of infected tubers is what is required and variations in the 
amount of disease on individual tubers are rarely of importance. Whether 
the tubers are in the soil or in the clamps the percentage of infection is 
easily obtained by the counting of samples, and if the samples are sufficient 
in number and taken at random, accurate estimations for the whole crop 
can be made. 


Moore, W. G. (1943). The measurement of plant diseases in the field. Trans. Brit. mycoL 
Soc. XXVI, i28--35. 

[Accepted for publication 15 March 1947) 

[ 142 ] 



A visit to Wheatfen Broad, Surlingham, Norfolk, from Friday to Monday, 
24-27 May 1946, marked the occasion of the first post-war Spring Foray. 

This visit was made possible by the kindness of Mr E. A. Ellis who, in 
addition to entertaining a party of members at his home at Wheatfen, 
placed his workroom and mycological library at the disposal of the visitors 
and gave unstintingly of his unrivalled knowledge of the locality. The 
Society is greatly indebted to Mr and Mrs Ellis for all they contributed to 
the success of the Foray. 

Wheatfen Broad is an area of some 150 acres of Yare Valley swamp 
situated about seven miles from Norwich, to the north of Rockland Broad, 
and bounded on the east by the river Yare and on the slightly higher ground 
to the south-west by Surlingham Wood. The main estate consists of a series 
of ‘marshes’ intersected by several miles of shallow streams connecting 
a chain of small broads navigable to a punt. Tidal communication with the 
Yare is responsible for a regular circulation of water and the normal rise 
and fall of 8 in. in the water level. The characteristic features of the 
vegetation are reed swamp, Glyceria marsh, Cladium fen, and sallow-ash 
carr. This area, which is of great biological interest, and which will it is 
hoped be permanently preserved, has been studied intensively for the past 
ten years by a group of naturalists who have published their results in 
a series of papers in the Transactions of the Norfolk and Norwich Naturalists 
Society (see Ellis, xni, 422-51, 1934, for a general account of the vegetation 
and XV, 191-219, 1941, for a list of micro-fungi by the same author). 

Most of the twenty-five members and friends attending the Foray found 
themselves on unfamiliar ground and all found much of interest. The main 
objective was micro-fungi and the collections made represented more than 
a hundred species belonging to diverse groups. All the finds have been 
entered on the card index kept by Mr Ellis with a view to the eventual 
publication of a critical account of the fungi of Wheatfen so that here it has 
only been thought necessary to supplement comments on some of the more 
interesting records with a list of fungi found in association with certain 
marsh plants. Specimens of most of the finds have been deposited in the 
herbaria of Mr E. A. Ellis, the Imperial Mycological Institute, or the 
Royal Botanic Gardens, Kew. 

Helicosporium phragmitis, vfhic]x is not included in Wakefield and Bisby’s 
list of British Hyphomycetes, was found in abundance on decaying stems of 
Glyceria maxima. These, according to A. E. Ellis, are its characteristic habitat 
in the Yare Valley but it has ^so been recorded on Phragmites, Phalaris 
arundinacea, zxl 6 l Calamagrostis arundinacea. Clasterosporium canicinum on Carex 
attracted attention, for this fungus (originally described by Schweinitz in 
North America) appears to have been recorded for Europe only from 

spring Foray ^ 1946 143 

Norfolk. Among other Hyphomycetes, Myrothecium inundatum Fr. on an old 
sporophore of Russula adusta and Pachybasium hamatum (Bon.) Sacc. found 
growing with Rhytisma acerinum on sycamore leaves may be mentioned, 
while Prof. C. T. Ingold listed seven aquatic species on dead leaves in the 
tidal ditches. The resemblance of Mitrula sclerotipus growing from sclerotia 
on blackened remains of meadowsweet leaves to species placed in Ver- 
patina Whetzel & Drayton, was noted and Pachydiscia marchantiae (Berk.) 
Bond, was found on the liverwort Conocephalum. The most noteworthy 
rust recorded was Puccinia persistens Plowr., forming abundant aecia on 
Thalictmm flavum. Larger basidiomycetes were infrequent but Miss Wake- 
field added six species of Cortichm^ including C. niveocremeum and C. 
roseocremeum^ and nine of Peniophora^ including P. Candida Lyman (syn. P. 
aegerita) with the Aegerita stage, P. leprosa^ and P. pub era to the list. There 
have been many records of entomogenous fungi from Wheatfen. During 
the Foray Gibellula aranearum (Schwein.) Syd. on spiders 2 aoA. Isaria farinosa 
(Holmsk.) Fr. on moth pupae were found. 

Acknowledgement must be made to Mr M. B. Ellis, Dr R. W. G. 
Dennis, Mr S. J. Hughes, and Mr E. W. Mason for making critical 

List of fungi associated with certain marsh plants 
AT Wheatfen Broad 

Caltha palustris. Ramularia calthae (Erikss.) Lindr. 

Carex, Marasmius menieri (on C. riparia)^ Arthrinium sporophlaeum Fr., 
Clasterosporium caricinum Schw., Gonatosporium puccinioides 
(Fr.) Corda, Volutella melaloma Berk. & Br. 

Cladium mariscus, Pistillaria aculeata, Tetraploa aristata Berk. & Br. 
Equisetum palustre. Stamnaria persooni (Fr.) Fuckel. 

Filipendula ulmaria, Mitrula sclerotipus Boud. 

Peronospora calotheca de Bary. 

Glyceria maxima. Gibberella zeae (Schw.) Petch, Coprinus (? urticaecola), 
Dematium hispidulum Fr., jHelicosporium phragmitis H5hnel. 
Myosotis. Synchytrium aureum Schroet., Erysiphe cichoracearum DC. 
Myrica gale. Ovularia destructiva (PhilL & Plowr.) Massee. 

Peucedanum palustre. Plasmopara nivea (Ung.) Schroet. (also on Aegopodium 
and Angelica). 

Phragmites communis. Dasyclypha controversa (Cooke) Rehm, Pyrenopeziza 
arundinacea (DC.) Boud., Lophodermium arundinaceum (Fr.) Ghev., 
Napicladium arundinaceum (Corda) Sacc. 

Poa trivialis. Passalora graminis (Fuckel) Holinel. 

Salix. Bertia moriformis (Fr.) de Not., Calyculosphaeria tristis (Fuckel) 
Fitzpat., Ghaetosphaeria phaeostroma (Dur. & Mont.) Fuckel, 
Diaporthe eres Nits., Eutypa flavovirens (Fr.) TuL, Gloniopsis 
levantica Rehm, Hypoxylon rubiginosum (Fr.) Fr., Lophiostoma 
salicum (Fabr.) Sacc., Nitschkia cupularis (Fr.) Karst., Rosellinia 
aquila (Fr.) de Not., Pholiota erinacea (Fr.) Quel., Goniothecium 
amentacearum Corda. 

[ 144 ] 


Flax Diseases. By R. McKay. (Flax Development Board, Ltd.; 2 Kildare 
Place, Dublin, 1947.) 55 pp., 52 figs. Price 5 r. 

At the beginning of the War flax jumped into prominence as an important crop: and in 
England and Wales alone the area devoted to it increased rapidly from about 4500 acres 
in 1939 to over 60,000 acres in 1944. Despite the wise precaution of ensuring that all 
flax seed was suitably disinfected with Nomersan before sowing, some anxiety was not 
unnaturally felt about the possible development and spread of disease in the crop, and 
this in turn revealed a gap in plant disease literature, for no general account of flax 
diseases was available to farmer or adviser. This gap has now been filled very adequately 
by Prof. McKay’s informative booklet, which provides all those interested in flax cultiva- 
tion with a concise account of its pathology and a means whereby all the significant 
diseases and pests of the crop can be recognized in the field. The text is a happy blend 
which satisfies the plant pathologist without diminishing the value of the bulletin as 
a guide for the interested factory fieldsman and grower. Well-selected references are given 
after the description of each disease, though a few recent ones that might be expected in 
a publication appearing in 1947 are missing. There is a glossary of technical terms, and 
indexes to common and scientific names as well as to authors. The bulletin is very 
attractively designed and profusely illustrated: it would be interesting, however, to see 
the effect of applying colour work to this rather difficult photographic subject. Myco- 
logists may well be puzzled by the invariable use of brackets for the authorities of the 
fungus names. 

W. C. Moore 


It is intended, as soon as circumstances allow, to issue the 
Proceedings of the Jubilee Celebrations of the British Mycological 
Society as Volume XXX of the Transactions’, this volume will be 
sent to all who were members on 30 June 1947. In accordance 
with this intention, the present volume is numbered XXXI. 

VoL XXXI, Parts III & IV 

Jime 1948 


; By a. a. PEARSON and R. W. G. DENNIS 

The present list of species will, we hope, be of service to students. It can 
be used both as a check list and as an indication of the modern aspects 
and tendencies in the taxonomy of this group of fungi. 

Our fungus books are cluttered with names that have little or no 
meaning, and it is time such names were withdrawn. This was the first 
purpose which induced us to undertake the task of preparing a list of 
species which we believe to be distinct and to have been correctly reported 
as British. 

The specific epithets which we have excluded are: 

^ (i) Synonyms. 

(2) Names attached to inadequate descriptions. 

(3) Species unlikely to occur in Britain. 

(4) Species which have been interpreted in more than one sense. 

(5) Errors due to carelessness. 

(6) Alien species included in existing floras on account of casual occur- 
rences among hothouse plants. 

■ The sifting was no easy task, and the result will not please everybody. 
Objections will come from two sides. We shall be told that we have been 
too drastic. That will be put right if and when species wrongly excluded 
are rediscovered and more fully described. Some will think we have left 
in species that are not really known to any living mycologist. That is true, 
but species may be so rare that only by very fortunate circumstances are 
they likely to be met with by a mycologist competent to name them. If 
they have points of distinction that make it likely that they are authentic 
species, different from any familiar to us, we have preferred not to discard 
them — ^at least for a few years more. 

It does not follow then that aU the specific names left in the list are 
guaranteed to be of permanent value. We are all too familiar with the 
Hght-hearted way in which names were given to fungi in the old days and 
the many pitfalls that beset mycologists when dealing with critical groups. 
All we can claim is that we have done our best to produce a list of species 
which actually do exist in Britain. 

The total number of the Agaricales left in our Ust is 1234, contrasted 
with about 1870 species contained in the latest authoritative work, Carleton 
Rea’s British Basi£orrgicetae^ and those subsequently recorded. The Boletales 
number 47 compared with 70. This does not take into acGount the varieties 
which we have dealt with somewhat relentlessly. About one- third of the 
names then have been excluded, and the reasons will be found in the 
explanatory notes at the end of each genus. 

VoL XXXI, Parts I & 11 : issued 1 1 September 1947. 



146 Transactions British Mycological Society 

There are, however, many more agarics that await determination. As 
we get to know the fungi more, the twofold process of Tumping’ and 
‘splitting’ must go on. The new methods of diagnosis, which lay greater 
stress on microscopic features and give some diagnostic value to chemical 
reactions, will certainly lead to the multiplicity of names. All we can 
hope for is that the tendency to split into iimumerable ‘Jordanian’ species 
will not be carried to excess. 

Our second object was to bring forward a transitional grouping which 
would bear some relation to modern tendencies in taxonomy and at the 
same time not interfere too violently with the Friesian framework which 
stiU seems the most practical. This has already been done in a narrower 
field widi the publication by the Essex Field Club of the List of the Fungi of 
Epping Forest, by A. A. Pearson (1938). We have adhered to the general 
classification based on the colour of the spores which stiU seems as good 
as any other, and is the most convenient for the display of specimens at 
fungus forays. In recent years, however, many new genera have been 
introduced, and we have indicated these in our sectional headings by 
mcbsing the new generic epithets in brackets. The new genera may thus become 
familiar to those who use this list, and later they can be adopted if thought 
worthy of generic rank. Much as we appreciate the activities of the 
modern mycologists who have been carving so many new genera out of 
the old, we think that many of these are based on such small difierences 
that they are scarcely worthy of more than sectional status. 

The works consulted are too numerous to mention. We have been 
fortunate in having open to us in the Herbarium of the Royal Botanic 
Gardens, Kew, the full range of books on systematic mycology. In the 
citation of authors we can hardly hope to have avoided occasional errors, 
and there are doubtless many specific names which will require changing 
when all the sifting has been done to bring them into line with the Inter- 
national Rules of Botanical Nomenclature. In a few cases, to avoid con- 
fusion, we have preferred not to disturb well-established epithets when 
there was some slight doubt as to the claims of priority. On the other 
hand, many familiar names have to go. These are mostly those which 
must be replaced by the epithets used by Fries in the Systema Mycologicum 
(1821), which, according to the International Rules, is the starting-point 
for the groups we have dealt with. The specific epithets m this work are 
valid when we clearly know what species he was referring to, even though 
in later books, he discarded them — as he often did. 

In our author citations, in most cases we have followed the usual 
practice in quoting Fries as the generic authority in instances in which he 
actually referred to sections or subgenera of Agarics. This does not comply 
strictly with the International Rules of Nomenclature, but we have felt 
unable to spare the time necessary for the exhaustive search of the literature 
required to establish who first raised the subgenus to full generic r ank in 
the case of each separate species. This sterile occupation appears particu- 
larly purposeless, as many names are likely to be changed afresh as soon 
as some of the sections of Friesian genera are accorded in Britain the 
generic status they have already attained elsewhere. 

Revised List of British Agarics and Boleti 147 

Neither have we followed exactly the Recommendation xxxii of the 
International Rules, but have continued what has hitherto been the 
common practice of citing the original author of a species in curved 
brackets instead of in square brackets or by using ex before the author 
whose description is accepted as defining the species. Some authors appear 
to have misunderstood this recommendation and place ex before the earlier 
author — Fr. ex BuU. — ^instead of before the later author — ^BulL ex Fr, 

A last note about the classification. It is only tentative and many of 
the sections are somewhat artificial. Some genera would be excluded from 
the Agaricales if due weight were given to the histological data accumu- 
lated in recent years. Doubtless there will be many changes in the taxo- 
nomy of this vast group of the fungi. With our present limited knowledge 
we have not felt called upon to do more than bring forward a transitional 
arrangement which, while not satisfying the more learned, will, we hope, 
be of service to mycologists who want an easily accessible outline classifi- 
cation of the British species. Some such critical revision of the published 
names is, in any case, necessary before a revision of the British Agaric 
Flora can be contemplated. 

(Acanthocystis), i6i 


(Deconica), 184 

(Lentinellus), 162 

Acetabularia, 173 

(Delicatula), 160 

Leutinus, 162 

(Agrocybe), 172 

(Dermocybe), 178 

(Lepidella), 148 

(Alnicola), 174 

Dictyolus, 168 

Lepiota, 149, 150 
(Lepiotella), 149 

Amanita, 148 

Drosophila, 183, 184 

Amanitopsis, 148 

(Lepista), 153 

(Androsacens), 159 

Eccilia, 171 

Leptonia, 170 

Anellaria, 185 

Entoloma, 170 

Leptotus, 168 

Armillaria, 150, 15 1 
'^Armiilariella), 150 

(Leucopaxillus), 153 

Flammula, 1 73 

(Leucocortinarius), 151 

^Aspropaxillus), 153 

(Fulvidula), 174 

(Limacella), 149 

(Astrosporina), 181 

(Limacium), 163 

Galera, 176 

Locellina, 173 

(Baeospora), 159 

(Galerina), 172, 174, 175, 176 

Bolbitius, 176 

(Glaucospora) 150 

(Macrocystis), 174 

Boletinus, 188 

Gomphidius, 188 

Marasmius, 158 

Boletus, 188 

Gyrodon, 188 

(Melanoleuca), 152 

Gyroporus, 188 

(Mucidula), 150, 157 

(Gamarophyllus), 163 

Mycena, 155 

(Gantharellula), 153 

Hebeloma, 172, 173 

(Mycenella), 155 

Cantharellus, 168 

(Hydrocybe), 179 

(Myxacium), 177 

Ghitonia, 186 

Glarkeinda, 186 

(Hygrocybe), 163 

(Hygrophoropsis), 153 

Naucoria, 172, 174 

Claudopus, 171 

Hygrophorus, 163 

(Nematoloma) , 1 83, 1 84 

Glitocybe, 153 

Hypholoma, 183, 184 

Neurophyllum, 168 

Glitopilus, 169 

Nolanea, 170 

Gollybia, 157 

Inocybe, 180 

Nyctalis, 162 

(Gonocybe), 172 

(Inoloma), 178 

Goprinus, 186 V 

(Ixocomus), 188 

Omphaiia, 160 

(Gortinellus), 151 

(Oudemansiella), 157 

Gortinarius, 177 

Graterellus, 168 

(Krombholzia), 189 

(PanaeoHna), 184 

Grepidotus, 176 

Laccaria, 154 

Panaeolus, 185 

Crinipellis, 159 

(Lacrimaria), 183 

(Panellus), 162 

(Gystoderma), 149 

Lactarius, 164 

Panus, 162 


iiS Transactions British Mycological Society 

Paxillusj 182 
(Phaeocoilybia), 174 
(Phaeolepiota), 172 
(Phaeomarasmius), 172 
Phaeopoms, 190 
(PMegmacium), 1 77 
Pboliota, 172 
Phylloporus, 188 
(Piyllotopsis), 176 
Pilosace, 186 
(Pleurotellus), 161 
Pleurotus, 161 
Plicatura, 168 
Pluteolus, 169 
Pluteus, 169 
Porphyrellus, 188 

Psalliota, 186 
(Psathyra), 184 
Psathyrella, 184 
(Pseudocoprinus) ,185 
Psilocybe, 184 

(Rhodocybe), 152 
(Rhodopaxillus), 152, 153 
(Rhodophyllus), 170 
(Rhodotus), 162 
Ripartites, 169 
(Rozites), 172 
Russula, 166 

Schizophyllum, 162 
(Schulzeria), 150 

Strobilomyces, 188 
Stropharia, 182 

(Tapinella), 182 
(Telamonia), 178 
(Togaria), 172 
Tricholoma, 15 1 
Trogia, 168 
Tubaria, 175 
(Tylopilus), 188 

Volvaria, 168 

(Xerocomus), 189 
(Xeromphalina), 159, 160 
Xerotus, 163 
(Xemla), 157 


Gen. AMANITA (Pers.) Fr. 


Amanita phalloides (VailL) Fr. 
verna (Bull.) Fr. , 
virosa Fr. 

Semi 4 imbatae Gilbert 
Amanita porphyria (A. & S.) Fr. 
citrina (SchaefF.) Roques 
* var. alba Price 
gemmata (Fr.) Gillet 
Eliae Quel. 

Floccosae Gilbert 
Amanita muscaria (Linn.) Fr. 
var. Formosa Fr. 

Amanita muscaria var. regalis Fr. 
solitaria (Bull.) Fr. 
pantherina (DC.) Fr. 

f. robusta Pearson 
excelsa Fr. 
rubescens (Pers.) Fr. 

var. annulo-sulphurea Gillet 
aspera (Fr.) Quel. 

{Lepidella Gilbert) 

Amanita Vittadini (Moretti) Vitt. 
echinocephala Vitt. 


Amanitopsis vaginata (Bull.) Roze Amanitopsis nivalis (SchaefF.) W. G. Sm. 

fulva (Grev.) Rea inaurata (Seer.) Boud. 

AiTianitd ucuhata Quel. — ^probably A. echinocephctla* 
cariosa IPr.— A, excelsa. 

Emilii BdQl.---SL form o£ A. miiscaria. 
luiea Otth.—doubtful species. 

(Batsch.) Fr. 1836=^. ci«n«a (Schaeff.) Roques, 1832. 
nima ¥r.‘--prohahly A, solitaria. 

ovoidea (BuU.) Fr.— a Mediterranean species reported as far north as Paris. There are two British 
records, neither very convincing. 
recutita ’Fr.= A. porphyria. 

spusa FT. Epi^. i^Q^A.exceUaEv. Syst. mi, There is little doubt that these two epithets 
represent the same species, A. excelsa hoing the form with the stem starting from deep in the 
^ound, which causes it to have a less characteristic obconic shape at the base. 
strobthformts A. solitaria. 

w (Lam., BuU., Pers.) Fr. h rather a problem. Fries in Syst. Myc. states ^sFpes. . . lac^ciim\ 

which pomts to what later he caUed .^^^^^^^^ However, he cites BuUiard, tab. 108, in 
de /a where the stem is perfectly smooth apart from the pendulous ring. To avoid 

Revised List of British Agarics and Boleti 149 

confusion it is better to use virosa for the Amanita with rough squamulose stem. Until recent 
years A, verna (Bull.) was looked upon as a variety of A. phalloides^ but is now accepted as 
a separate species. 

Amanita Vittadinii (Moretto) Vitt. — ^left in, but doubtful if found in northern Europe. Gke. i//. 

33 (3^} copied from Krombholz. 

Amanitopsis strangulata (Fr.) Roze—^. inaurata, 
adnata {y^,G,Sm.) Sacc,=i Amanita gemmata. 


{Limacella 'Eavle em. yLaixe) 
Lepiota illinita Fr. 
guttata (Pers.) Fr. 
glioderraa Fr. 
arida Fr. 
delicata Fr. 

{Lepiotella Gilbert) 

Lepiota irrorata Quel, 
medullata Fr. 

Procerae Fr. 

Lepiota procera (Scop.) Fr. 
mastoidea Fr. 
excoriata (Schaeff.) Fr. 
gracilenta (Klrombh.) Fr. 
rhacodes (Vitt.) Fr. 

var. puellaris Fr. 
nympharum Kalch. 
permixta Barla 
Olivieri Barla 
Badhamii B. & Br. 
naucina Fr. 
holosericea Fr. 

Echinatae Fayod 

Lepiota acutesquamosa (Weinm.) Fr. 
echinella Quel, 
hispida (Lasch) Fr. 

Granulosae Yr, 

{Cystoderma Fayod) 

Lepiota granulosa (Batsch) Fr. sensu Lange 
cinnabarina (A. & S.) Fr. 
amianthina (Scop.) Fr. 
carcharias (Pers.) Fr. 
haematites (B. & Br.) comb. nov. 
lilacea Bres. 
nigromarginata Mass. 

Micaceae Lange 
Lepiota Bucknallii B. & Br. 
sistrata Fr. 
rosea Rea 

Pilosellae Kiihner 

Lepiota Georginae W. G. Sm. 

Anomalae Lange 

Lepiota Eyrei (Mass.) Lange 
haematosperma (Bull.) Bond. 

Fusisporae Lange 

Lepiota clypeolaria (Bull.) Fr. 
laevigata Lange 
alba (Bres.) Sacc. 
erminea Fr. 

Stenosporae Lange 

Lepiota fulvella Rea 
castanea Quel, 
cristata (A. & S.) Fr. 
pseudo-felina Lange 
Grange! (Eyre) Lange 

Ovisporae Lange 

Lepiota constricta (Fr.) Quel, 
meleagris (Sow.) Fr. 
bnmneo-incarnata Chod. & Mart, 
helveola Bres. 

fusco-vinacea Moller & Lange 
citrophylla B. & Br. 
felina (Pers.) Fr. 
gracilis (Quel.) Rea 
scobinella (Fr.) Quel. & Bat. 
clypeolarioides Rea 
serena Fr. sensu Kiihner 
parvannulata (Lasch) Fr. 

Leucobolbitiits Lange 

Lepiota cepaestipes (Sow.) Fr. 
lutea (Bolt.) Quel. 

Brebissonii Godey 

150 Transactions British Mycological Society 


{Armllariella Karst.) {Mucidula Pat.) 

Armillaria melJea (Vahl.) Fr. Armillaria mucida (Schrad.) Fr. 


Lepiota amimthina (Scop.) Fr. — ^it would perhaps be more correct to name this species L. granulosa^ 
since Fries used it in SysU Myc. citing X. amianthina Scop, as synonym; but he also cited 
Bulliard’s two plates of Agaricus ochraceus which appear to represent both species in question. 
arida Fr. — ^left in, but the record reqmres confirmation. 
atro-crocea W.G.Sm. — ^from Somerset (1903), description inadequate. 
biornata B. & Br. — alien, only in hothouses. 
cepaestipes YQx. cretacea GTev.=:L. BrebissoniL 

consiricta (Fr.) Qud. — probably the same as Tricholoma leucocephalum but left as separate species 
for the present. If proved to be identical, the former epithet would be valid. 
emplmtrim Ckc, ^ rkacodes. 

Friesii {Ju2cs,ch) 'Fx,'=^L. acutesqmmosa. 
ianthina Gke. — an alien; in stove at Kew. 
lentieularis Ijzsch— L. guttata* 
martialis^ Gke. & Mass.— alien, on pahn stem. 
mesomorpha (Bull.) Fr. — doubtful record. 

B. & Br.— a Geylon species, in greenhouse. 

Pauletii'Px. — doubtful species. 

polysticta Berk, (i860) — both description and figure in Gke. ///. 41 (30) suggest X. granulosa, 
pratensis (Bv.) 'Kez.^L. clypeolaria. 
pseudo 4 icmophora 'R.e2t=^ L. lutea. 

rosea Rea — left in list, but Lange thinb it to be X. rufescensB. & Br, 
seminuda (Lasch) "Ex.—L. sistrata. 

submarasmoides Speg. — the original diagnosis refers to a minute agaric found in the Argentine 
which hardly corresponds to the species described by Rea. 

Glaucospora Rea — established to include species of Lepiota with spores tinted blue or green, 
a character which hardly calls for a new genus. Glaucospora Eyrei has been listed as Lepiota 

Schuk&ria Bres.— this genus has also been withdrawn. The two species in the British list both 
suggest the presence of a veil though not forming.a ring on the stem; but this happens with 
many species oi Lepiota. Schulzeria Grangei Eyre takes its place in the group of Lepiota with 
projectile-shaped spurred spores, ecnd Schulzeria lycoperdoides Gke. & Mass, is doubtful. 
Armillaria Fr. — the heterogeneous collection of agarics attached to this genus had only one 
feature in connnon: the presence of a ring. Fries himself from the Syst. Myc. onwards 
divided up his section Armillaria into groups which were respectively akin to Tricholoma^ 
Clitocybe, Pleurotus and Collybia. In the present list we have followed the example of modern 
authors and transferred the appropriate species to these different genera. Only two species 
both common, are left to represent Armillaria. Even these have been abandoned hy some 
authors for other genera. 

Armillaria bulbigera {h. 81 Er:) Ex. => Tricholoma bulbigera. 
caligata {yiv.) Ex.^ Tricholoma caltgata. 

(Inzenga) Fr.— a form of A 
colossa (Fr.) Boud. = Triclwloma colossum, 
constricta Ex.=^ Lepiota cons^icta. 
delicata (JEx.) Bond.^ Lepiota delicata. 
denigrataEx. — ^xohQhly Pholiota erebia. 
focalis Ex.— Tricholoma robustwn. 
haematites B. Sc Bx.— Lepiota haematites. 

Jasords Cke. Sc Mass. Cooke’s figure, PL 1113 (955}, shows a robust form of Lepiota 

Revised List of British Agarics and Boleti 151 

Armiltaria mellea (Vahl) Fr. — this polymorphous species has a large number of varieties, none 
of which is included in this list. The var. tabescens is accepted as a separate species and 
will be found under Clitocybe. C. ectypa is also related but the British record is doubtful. 
robusta {A. S.) 'Fx,^Tricholoma robustum. 

ramentacea (Bull.) Fr. = Tricholoma ramentacea. 

tufa (Batt.) Quel.—Gooke’s figure 51 (33) represents Tricholoma robustum, md there has been 
no other collection published. 

subcam (Schum.) Fr.- — doubtful; Cooke’s figure 57 (47) is Lepiota Brebissoniu 



Tricholoma gambosum Fr. 
album (Schaeff.) Fr. 
lascivum (Fr.) Gillet 
spermaticum Fr. 
resplendens Fr. 
columbetta Fr. 
sulphurescens Bres. 
inamoenum Fr. 
leucocephaium Fr. 

(See also Clitocybe connata) 


Tricholoma aggregatum (Schaefif.) Seer, 
pergamenum (Cke.) comb. nov. 
cartilagineum Fr. non Bull, 
cinerascens (Bull.) Fr. 
loricatum Fr. 
immundum Berk, 
semitale (Fr.) Ricken 
infumatum (Bres.) comb. nov. 
leucophaeatum Karst. 


(a) Limacina Fr. 

Tricholoma albo-brunneum (Pers.) Fr. 
ustale Fr. 
pessundatum Fr. 
fulvum Fr. 
acerbum (Bull.) Fr. 

(b) Flocculosa Fx, 

Tricholoma imbricatum Fr. 
vaceinum (Pers.) Fr. 
psammopus (Kalch.) Fr. 
macrorhizum (Lasch) Fr. 

(r) {Armillaria'Fr,) 

Tricholoma robustum (A. & S.) Fr. 
caligatum (Viv.) Fr. 
colossus Fr. 

aurantium (Schaefif.) Fr. 


Tricholoma flavo-virens Fr. 

( = equestre (Linn.) Fr.) 
fucatum (Fr.) Gillet 
sejunctum (Sow.) Fr. 

var. corypheum (Fr.) comb. nov. 
portentosum Fr. 
sulphureum (Bull.) Fr. 

var. bufonium (Pers.) Fr. 
saponaceum Fr. 

var. squamosum Cke. 
rutilans (Schaeff.) Fr. 

var. variegatum (Scop.) Fr. 
decorum Fr. 

{Armillaria Fr., Cortinellus Roze and 
Leucocortinarius Lange) 
bulbigerum (A. & S.) Fr. 


Tricholoma terreum Fr. 
scalpturatum Fr. 

Inocybeoides Pearson 
triste (Scop.) Fr. 
atro-squamosum (Ghdv.) Sacc. 

var. squarrulosum Bres. 
horribile Rea 
orirubens Quel, 

var. guttatum (Schaeff.) comb. nov. 
inodermeum Fr, 
virgatum Fr. 
gausapatum Fr. 

{Armillaria Fr.) 

Tricholoma ramentaceum (Bull.) Qjuel. 


Tricholoma carneum (Bull.) Fr. 
onychinum Fr. 
ionides (Bull.) Fr. 

var. obscurissima Pearson 
var. persicolor (Fr.) Bres. 

Transactions British Mycological Society 


Dermdoma Lange 

Triciioioma atrocinereum (Pers.) Fr. 
cuneifoiium Fr. 

{Melanoleuca Pat.) 

Tricholoma melaleucum (Pers.) Fr. 
var. polioleucum (Fr.) Gillet 
var. adstringens (Pers.) Quel, 
grammopodium (Bull.) Fr. 
cognatum Fr. 
evenosum Sacc. 
turritum Fr. 

(Ehodopaxillus Maire) 
Trichoioma nudum Fr. 

var. lilaceum (Quel.) Kiihner & Maire 
glaucocanum Bres. 
sordidum Fr. 
personatum Fr. 
panaeolum Fr. 
truncatum (SchaefF.) Quel, 
irinum Fr. 

(See also Clitocybe) 

(Bhodocybe Maire) 

Trichoioma caelatum Fr. 


Trichoioma albellum Fr. ^gambosum Fr. 
amplum {?t,Ts,) 'R.tdi.= T aggregatum^ 
amarim {K. ^ Clitocybe dmaraVv^ 

amethystinum {Scop,) personatum 'Pv^ 

amicum Fr. — doubtful species. 
arcuatum (Bull.) Quel. = T cognatum Fr. 
argyraceum (Bull.) Fr. = T. scalpturatum Fr. 
horealePt.— T, irinum Px, 
brevipes {PviPi) Px,'==^T.rnelaleucumioxm, 
ceniurio Kalch. — doubtful. 
cerinum (Pers.) Fr. — doubtful record. 
chrysites (Jungh.) Gillet = T scalpturatum Fr. 
circumtectum Ck.t. 7 =T, atro^squamosum* 
civile Px.== Clitocybe amaraPx, 

cnista Px,=see, Lange form of saponaceum, but most authors attach it to Melanoleuca group. 
compactum Fr. — added to British list by Massee in 191 1 without substantiating data. 
duracinum Ckc. ^cinerascens, 
elytroides (Scop.) Fr. — doubtful. 

equestre (Linn.) Px,=flavo-virens as in Syst, Myc.^ though Fries abandoned this later for the 
Linnean name. 
exscissumPx.=^T. melaleucum. 
fallax Peck . — doubtful record. 
flavobnmneum Px.=:T.fulvwn (DC.) Pr, 

focale Pxj ^record uncertain. Gke. III. 49 (245) suggests a form of T. aurantium. 
focale var. Goliath Fr. — also uncertain, 
fulvellumPx.= T.fulvum. 

Georgii (Glus) Fr.= T. gambosum. 
hordum Px . — doubtful species. 

hcrribile P.eB, ^left in, but doubtful if other than a form of orirubens. 
kumile (Pers.) Fr.= T’. melaleucum. 
interveniens K.zxst . — ^no British record. 
irregulareK. 3 xst.^albobrunneum. 

ImoiumPx. melaleucum. 
luridum (Sch.2ie^.) Px.^ T. saponaceum. 
luteociMnumP.eQ.==^Hygrophorus ImorumPialch.. 

(Lasch) Fr.— identity doubtful. 
munnaceum (Bull.) Pt.^T. virgatum. 

myomyces Fr.— this epithet was used in Syst. Myc. for the whole group of Trichoioma of which 
r. ^am IS the commonest species. If the rules are strictly foUowed T. myomces should 
perhaps replace terrewn. We prefer to use the epithet as a Section heading. 
nktitans Pi. =^ T.fulvtm. ° 

Revised List of British Agarics and Boleti 153 

Tricholoma opicum Fr. — ^identity uncertain. 
opiparum QKkB.,^Hygrophorus nmonus. 
paedidum Fr. = T, melaleucum, 
patulum Fr. = T. melaleucum. 
pes-caprae Fr. == T. cinerascens. 
phaeopodium (Bull.) Quel. = T. melaleucum. 

porphyroleucum (Bull,) Fr. = T. melaleucum or T. ionides var. obscurissima. 
putidum^T. — Collybiaputida (Fr.) iCuhner & Maire. 
quinquepartitum^x . — ^uncertain species; probably T. 
saevum Gillet= T. personatum Fr. 

Schumacheri Fr. — ^identity doubtful. 

Stans Fr. = T. pessundatum. 
subpulverulentum (Pers.) Fr. = T. melaleucum. 
sudum Fr. — doubtful record. 
tenuiceps Gke. & Mass. = CoUyhia platyphylla. 
tozWwm (Pers.) Fr.— doubtful record. 
turritum Fr. — ^probably T. melaleucum. 
unguentatum’Fx . — ^identity uncertain. 


Clitocybe clavipes (Pers.) Fr. 
obscurata Gke. 
inornata (Sow.) Fr. 

var. exilis Pearson 
odora (Bull.) Fr. 

var. Trogii (Fr.) Maire 
pseudoconglobata Rea 
tabescens (Scop.) Bres. 

{Aspropaxillus Kiihner & Maire) 
Clitocybe gigantea (Sow.) Fr. 

{Leucopaxillus Boursier) 

Clitocybe amara Fr. 
paradoxa Const. & Duf. 

{Cantharellula Singer) 

Clitocybe umbonata Fr. 
cyathiformis (Bull.) Fr. 
obbata Fr. 

{Lepista W.G.Sm. em. Pat.) 

Clitocybe flaccida (Sow.) Fr. 
gilva (Pers.) Fr. 

{Rhodopaxillus Maire) 

Clitocybe popinalis (Fr.) Bres. 
f. senilis Fr. 

mundula (Lasch) comb, nov. 

{Hygrophoropsis (Schroet.) Maire) 

Clitocybe aurantiaca (Wulf.) Studer 
var. albida Gillet 
var. nigripes Pers. 
albida (Fr.) Konrad 

Camosae Lange 

Clitocybe geotropa (Bull.) Fr. 
var. maxima (Fr.) Nuesch 
nebularis (Batsch) Fr. 


Clitocybe infundibuliformis (SchaefF.) Fr. 
var. catinus Fr. 
incilis Fr. 
sinopica Fr. 
paropsis Fr, 
vermicularis Fr. 
parilis Fr. 
trullaeformis Fr. 
himeola Fr. 


Clitocybe cerussata Fr. 
phyllophila Fr. 
monstrosa (Sow.) Gillet ’ 
pithyophila Fr. 
rivulosa (Pers.) Fr. 
dealbata (Sow.) Fr. 
candicaiis (Pers.) Fr. 
ericetorum (Bull.) Fr, 
connata (Sebum.) Fr. 
tuba Fr. 

154 Transactions British Mycological Society 

Hygrophame, Decurrentes 

Giitocybe vibecina Fr. 
expalkns (Pers.) Fr. 
concava (Scop.) Fr. 
brumalis Fr. 
fritilliformis (Lasch) Fr. 
angustissima (Lasch) Fr. 
incana Quel. 


Giitocybe diatreta Fr. 
ditopus Fr. 
metachroa Fr. 
fragrans (Sow.) Fr. 
obsoleta (Batsch) Fr. 
albocinerea Rea 

Gen. LACCARIA B. & Br. 

Laccaria laccata (Scop.) Gke. 
proxima Boud. 
amethystina (Vaill.) Gke. 

Laccaria tortilis (Bolt.) Gke, 
nana Massee 
bella (Pers.) Gke. 


Giitocybe aggregata Tncholoma aggregata, 

amarella (Pers.) Fr. — ? form of C. {Rhodopaxillus) popimlis, 
cartilaginea (Bull, non Fr.) Bres. = Tricholoma loricatum. 
comitalis (Pers.) Fr. — ^identity doubtful. 
conglobata iyitt,) Tricholoma aggregatum. 
cryptarum (Letell.) B. & Br. — ^identity uncertain. 
curiipes Fr. — ^may be Tricholoma panaeolum, 

(yanophaea Fr. — pvohsbly Tricholoma nudum. 

decastes Fr.— in Syst. Myc. appears to be the same as Tricholoma cinerascens Bull., and in later 
'wotk^ oi Frits hzxdlY distmct from Tricholoma aggregatum. 
ectypa Fr. — ^British records are doubtful. Gke. III. 177 (126) is C. cyathiformis. 
elixa (Sow.) Berk. = C. inornata or C. Alexandra, 
gallinacea {Scop.) Fr.^ Omphalia hydrogramma. 

gangraenosa Fr. = Tricholoma leucophaeatum Karst. The epithet gangraenosa would have priority 
but its identity is in doubt. 

geotropa pBvXl.) Fr. Epicr . — ^we have hesitated whether to use the epithet gibba Fr. for this 
species. It is the Agaricus gibbus (Pers.) Fr. var. major Fr. Syst. Myc., but the type oigibbus is 

(Brond.) Rea — suggests Hygropkorus lucorum. 
inversa (Scop.) Er. — C.flaccida (Sow.) Fr. 

^^^<2 Fr. — ? Tricholoma panaeolum. 

(Sow.) Gill.^ — left in list but suggests C. 
obbata Gke. — ^remains in list only on authority of Gke. III. 168 (230). 

occulta Gk .^. — a Tricholoma of the Myomyces group. May be a good species but requires clearer 

opaca iyiith.) Fr. = C. cerussata. 
orbiformis Fr. — ^not clearly defined, 


pergamem Cke.= Tricholoma pergamenum. Suggests T. aggregatum but for the elliptical spores. 
polia Fr. — IC. inornata. 

(Lasch) Fr. — doubtful record. 
senilis Fr. =^hrra oi C. popinalis. 

socialis Fr.— description inadequate but Gke. III. 132 (134) looks distinctive. 
splendens iJF trs.) Fr.^ C. gilvaFr. 

suaveolens (Schum.) Fr, — though this may be a good species, it requires clearer definition before 
it can be considered distinct from C. fragrans. 
subalutacea (Batsch) Fr.— doubtful record. 
sub decastes Gke. & Tricholoma cinerascens. 

tornataFT, = C. cerussata. 

TrogiiFr. — Yax. oi C. odora. 

tumulosa (Kalch.) Fr.= Tricholoma aggregatum. 

Revised List of British Agarics and Boleti 155 

Clitocyhe venustissma'Fx. — Omphalia demissa, 
vemkosa Fr.— probably C. flaccida, 
midis (With.) Fr. = C. odora. 
zygophylla Cke. & Mass. = C. inomata, 

Laccaria Bella (Pers.) B. & Br. — ^left in list though doubtful, but the foetid smell would distinguish it. 
nana Massee — ^retained in list on account of the large spore 15-16/4, but requires confirmation. 

Gen. MYCENA Fr. 

Mycena pelliculosa Fr. 
vulgaris (Pers.) Fr. 

Roridae Kiihner 
Mycena rorida (Scop.) Fr. 

Alcalinae Konr. & Maubl. 
Mycena alcalina Fr. 
ammoniaca Fr. 
aetites Fr. 

(See also rubromarginata) 


Mycena pelianthina Fr. 
avenacea Fr. 
atro-marginata Fr. 
rubro-marginata Fr. 
eiegans Fr. 
rosella Fr, 
pterigena Fr. 
atro-virens Rea 
Seynii Quel. 

(See also haematopus) 

Lactipedes Fr, 

Mycena galopus (Pers.) Fr. 
var. alba FL Dan, 
var. nigra FL Dan, 
sanguinolenta (A. & S.) Fr. 
haematopus (Pers.) Fr. 

var. marginata Lange 
crocata (Schrad.) Fr. 

Basipedes Fr. 

Mycena stylobates (Pers.) Fr. 
bulbosa Gejp 
tenerrima Berk. 

(See also pterigena) 


Mycena amicta Fr. 


Mycena pura (Pers.) Fr. 
var. rosea Schum. 
var. multicolor Bres. 
var. alba Gillet 
pseudo-pura Cke. 
zephirus Fr. 
flavipes Quel. 

Adonis (Bull.) Fr. 
acicula (Schaeff.) Fr. 
atrocyanea (Batsch) Fr, 
urania Fr. 

Glutinipedes Fr. 

Mycena epipterygia (Scop.) Fr. 
viscosa (Seer.) Maire 
epipterygiodes Pearson 


Mycena polygramma (Bull.) Fr. 
vitilis Fr. non Kiihner 
erubescens V. Hohnel 
subalpina v. Hohnel 
galericulata (Scop.) Fr. 
inclinata Fr. 


Mycena lactea (Pers.) Fr. 
gypsea Fr. 
speirea Fr. 

hiemalis (Osbeck) Fr, 
fiavo-alba Fr. 
trachyspora Rea 
quisquiliaris (Joss.) Kiihner 

Granulatae Lange 
Mycena metata Fr. 
chlorantha Fr. 

var. pallida Pearson 
arcangeiiana Bres. var. 

Oortiana Kiihner 
filopes (Bull.) Fr. non Kiihner 
cinerella Karst, 
uracea Pearson 
fagetorum Fr. 
corticola (Pers.) Fr. 
mirata Peck sensu Smith 
capillaris (Schum.) Fr. 
debilis Fr. 
mucor (Batsch) Fr. 
polyadelpha (Lasch ex Fr.) Kiihner 
osmundicola Lange 


Mycena lasiosperma Bres. 


Transactions British Mycological Society 


JWyrma a/ro-a/Aa (Bolt.) Fr.— identity not clear, . , , , 

Fr.— placed next to M, cmmoniaca to which it approximates, but has no smell 

at£raMio~margimtal^T,=M, elegans^ 

Berkel^i Mass.-based on Cooke’s figure 224 (148) of M. excisa which represents a dark- 
coloured M, gaUriculata; but if Massee’s spores 5 X sJ-jLi are correct, it is certainly distinct. 
carneo-sanguinea Rea— cystidia not given but almost certainly a form of M, pelianthina. 
chelidonia Fr,— doubtful; may be a form of M. crocata. 
cinerea Mass. & Cross!. — ? form of aetites, 

citrinella (Pers.) Fr.— a doubtful species. Ricken records it for Germany, but his spores and 
cystidia suggest Jlavo-alba, 

clavicularis Fr.— variously interpreted; excluded for further observation. 
clavus (Linn.) Rea=Af. Adonis, 
coccima (Sow.) Quel.^M. Adonis, 
codoniceps Cke.— description inadequate. 

Fr.— identity uncertain. 

comimilis Oke.-— ni. ii ^0 (iiSQ) is tYpicsX M. ammoniaca. 
cruenia'Fx.^M. haematopus. 

debilis Fr. — ^left in list, but wants clearer definition. 
dilatata Fr. — ^probably M. stylobates, 
discopuslev. — piohzbly M. stylobates. 

dissiliens Fr.— has diJfierent interpretations; we do not know to which the British record refers. 
excisa (Lasch) Gillet.— probably M. galericulata. 
farrea (Lasch) Fr. — ^may be Lepiota sistrata. 
felleal^n%e—M. erubescensv.Wbhnel 
fusco^purpurea (Lasch) Cke. — uncertain. 
galericulata vax. caloptis 'Er.=M. inclinata. 

Iris Berk. = M. amictaFr. 

juncicola Fr. — description inadequate but has distinctive characters. 
leptocephala (^ers.) Ft. — seems identical with iW. c/ta/zwa. 

lineata (Bull.) Fr. — authors take different view^ about this. It is perhaps best to consider it 
2i y>2le form oi M. chlorantha. A.A.P. 
luteo~alb a (JSolt.) Fr. — ^identity doubtful. 
marginella Fr.— Cke. III. 1148 (957) appears to be M. amicta. 
nivea Q\x€l.=^M. gypsea. 

olivaceo-marginata Mass.—M. avenacea Fr. The smaller spores given by Massee are presumably 
an error. 

Fr.— has many interpretations. Best left out. 
paupercula Berk. — ^no microscopic details given. 
peltata Ft. — ^identity uncertain. 
plicato-crenata Fr.— hardly distinct firom M. epipterygia. 

plicosa Fr.— variously interpreted; withdrawn until more fully described from British specimens. 
plumbeaFr. — doubtful. 

prolifera (Sow.) Fr. — Sowerby’s figure t. 169 suggests M. inclinata. 

psammicola B. & Br. — ^not clearly defined. Alex. Smith describes it from American specimens 
as like a Galera. 

pseudo‘pura Cke. — ^usually considered to be a form of M. pura, but restored to specific rank by 
Kiihuer who states that the spores are non-amyloid. We have not been able to verify this 
as it appeails to be a rare species in Britain, M. pura has amyloid spores, 
jbwtofl Berk. & Cke.— uncertain. 
pura var. carnea Rea=5var. rosea Schum. 

rubro-marginata Fr.— we interpret this as being the Mycena which grows in vast numbers under 
conifers, with much the same smell as M. ammoniaca and M. alcalina. Frie^ in his Monographia 
says *est inter vulgatissimas Mycenasl It is true that he calls it ^ inodorus^ , but his sense of smell 
was defective, and sometimes the smell is weak or absent. Kiihner describes the species 
under M. capillartpesy Peck and Lange as M. plicosa var. marginata. 
rugosa Fr. — ^a mgose form of Af. 


Remsed List of British Agarics and Boleti 

saccharifera B. & Br. — ^probably M, tenerrima, 
setosa = Mmor, 

Qu61.— usually considered as confined to southern Europe, but has been recorded in 
Holland, so our one British record may be correct. 

Simillima Karst.— identity uncertain. 
stamea'^T* — ^probably a form of Af. when dry. 

strobilina Ft. = M. Adonis, 
sudora — doubtful species. 
fowMiy (Bolt.) Fr. — ^not clearly defined. 
tenella= M. metata. 

tintinnabulum Fr. — ^has many interpretations; not known to which the British record refers. 
irachyspora Rea — a good species, but the spores are smooth. Specimens from the trunk on 
which the original material grew are in the Kew herbarium, Royal Botanic Gardens. 
Unfortunately the name cannot be changed. 

mania Fr. — ^left in list, but this striking species awaits further study. The epithet has been 
adopted by Alex. Smith for an American species with warted cystidia. Its occurrence as 
British rests solely on Massee’s authority and requires confirmation. 
vitred Fr. — ^variously interpreted and best withdrawn. Cke. III. 237 (160) is probably M. 


and Striaepedes Fr. 

Gollybia platyphylla (Pers.) Fr. 
fusipes (Bull.) Berk, 
maculata (A. & S.) Fr. 
distorta Fr. 
butyracea (Bull.) Fr. 
fioccipes Fr. 
lacerata (Lasch) Berk, 
eriocephala Rea 


Gollybia rancida Fr. 
protracta Fr. 

putida (Fr.) Konr. & Maubl. 
inolens Fr. 
mephitica Fr. 
coracina Fr. 
murina (Batsch) Fr. 
misera (Fr.) Lange non Bres. 
atrata Fr. 
ambusta Fr. 
clusilis Fr. 

pseudo-clusilis Joss. & Konr. 
tylicolor Fr. 
tesquorum (Fr.) Bres. 
palustris (Peck) 

Collybia acervata FT. = Marasmius aceroatus. 
caldarii Berk. — ^an alien. 

cirrata var. Cookei Bres. — recorded by Bresadola in Icon. Myc. (1928), tab. 206, and based on 
Gke. ///. 197 (144), lower figure. It is cirrata with a yellow sclerotium; the type has none. 
(Scop.) Fr.— left in list but requires confirmation. 
confluensFr. — Marasmiusconfluens. 

{Oudemansiella Speg. (1882) 
Mucidula Pat. (1887)) 

Gollybia radicata (Rehl) Berk. 

{Xerula Maire) 

Gollybia longipes (Bull.) Berk, 

Laevipedes Fr. 

Gollybia nitellina Fr. 
extuberans Fr. 
collina (Scop.) Fr. 
succinea Fr. 
nummularia Fr. 
thelephora Cke. & Mass. 

Vestipedes Fr. 

Gollybia velutipes (Gurt.) Fr. 
f. rubescens Cke. 
f. lactea Quel, 
cirrata (Schum.) Fr. 
yar. Cookei Bres. 
var. ocellata Fr. 
tuberosa (Bull.) Fr. 
racemosa (Pers.) Fr. 
Stevensonii B. & Br, 

158 Transactions British Mycological Society 

Cottybia conigena 'Bt, — stt Marasmius noXts. 
crassifolia (Berk.) Bres. = 'Tricholomd iminundum Berk. 

Dorotfmae Berk. — an alien. 

dryophiU (^\iSS,)'^t.^Marasmms dryophU^ 

eriocephah Rea — ^may only be a form of C.fusipes but left in for further observation. 
esmlmta Vt.—Marasmius esculenfus. % 

eustygia Cke. = Tricholoma immundum. 

floccipes Fr. — Cke. III. 1 142 (i 168) looks authentic enough to be kept in the list but confirmation 
is needed. 

fodiens K^lch. — ? form ol maculata. 

fumosa (Peis.) Q}x€l. non Ft. = Tricholoma immundum, 

Henrieitae W. G. Sm. — probably a form of C. radicata. 
infunmta (Bves.) Rtsi—Tricholorna infumatum. 
lancipes Fr. — ^slender form of C.fusipes. 

laxipes (Batt) Fr. — ^identity uncertain. Cke. III. 191 (184) is probably Marasmius undatus. 
leucomyosotis Cke. & Smith (iBQ^)=Mycena palustris Peck (1872), recorded in this list 

luteifolia Gillct---~prohably Marasmius dryopkilics v2LT.funicularis. 

macilenta Fr. — ^record doubtful. Cke. III. 208 (268) represents Marasmius dryophilus var. funicu- 

mimica VI. G.Sm. — prohshly Naucoria cucumis. 
muscigena (Schum.) Fr. 1 —Mycma gypsea, 

ntmmidaria Fr. — ^left in list, but doubtful whether the British record is other than Marasmius 

ozesFv. — doubtful record. 

planipes (Brig.) Fr.— ? form of C. radicata. 

plexipesFr. — ?—C. protracta. 

prolbca (FI. Dan.) Fr. — can hardly be distinct from C. distorfa. 

psathyroides Cke.— needs confirmation; probably a Mycena but may be a form of C. radicata. 
pulla (Schaeff.) Fr. — ^may be a form of C. butyracea. 

Bres.— doubtful record. 
semitalis Ft. ^Trkkoloma semitale. 

Stevensonii B. & Br. — ^left in list but not known to modern authors. 
stridula (Schaeff.) Fr. = Tricholoma melalmcum, 

tesquorum Fr.— identity doubtful. It was identified by Lange with the rough-spored C. tylicolor, 
but later he preferred to associate the epithet with a smooth-spored agaric as Bresadola 
already had done. It is left in the list with this interpretation. 
ventricosa (BviH.) Ft. loxm of C. radicata. 
xanthopus Ft. = 7 M. dryophilus. 
xylophila (Weinm.) Fr. — ^may be Mycena galericulata. 


{a) Inodori 

Marasmius peronatus (Bolt.) Fr. 
fusco-purpureus (Pers.) Fr. 
acervatus (Fr.) comb. nov. 
lupuletorum (Weinm.) Fr. 
ceratopus (Pers.) Quel, 
confluens (Pers.) Karst, 
pruinatus Rea 
obtusifolius Rea 
oreades (Bolt.) Fr. 
oreadoides (Pass.) Fr. 

Marasmius Wynnei B. & Br. 
dryophilus (Bull.) Karst, 
var. funicuiaris (Fr.) Rea 
var. aquosus (Fr.) Rea 
exculptus (Fr.) Rea 
undatus Berk. 

(b) Alliodori 

Marasmius porreus (Pers.) Fr. 
prasiosmus Fr. 
scorodonius Fr. 
alliaceus (Jaeq.) Fr. 

Remsed List of British Agarics and Boleti 159 

(c) Ingrati 

Marasmius foetidus (Sow.) Fr. 
impudicus Fr. 
perforans Fr. 

{d) Conigeni 

Marasmius esculentus (Wulf ex Fr.) 

{e) {Baeospom Singer) 
Marasmius myosurus (Wulf ex Fr.) 

(a) Ramealini 

Marasmius ramealis (Bull.) Fr. 
Vaillantii (Pers.) Fr. 

Marasmius Menieri Boud. 
tricolor (A. & S.) Fr. 
amadelphus (Bull.) Fr. 

{b) {Androsaceus (Pers.) Pat.) 

Marasmius rotula (Scop.) Fr. 
graminum Lib. 
obtusifolius Rea 
androsaceus (Linn.) Fr. 
splachnoides Fr. . 

epiphyllus (Pers.) Fr, 
epiphylloides (Rea) Sacc. & Trotter 
Hudsonii (Pers.) Fr. 

(r) {Xeromphalina Kuhner & Maire) 
Marasmius cauticinalis (With.) Fr. 

Crinipellis stipitarius (Fr.) Pat. 


Marasmius actinophorus B. & Br. — ^may be M, graminum. 
amadelphus — ^left in, but confirmation needed. 

angulatus (Batsch) B. & Br. — probably M. VaillantiL 
archyropus (Pers.) Fr. — doubtful identity. 
calopus (Pers.) Fr. = Af. scorodonius, inodorous form. 
candidus (Bolt.) Fr.ssAf. VaillantiL 
caulicinalis QxL€l.^Crinipellis stipitarius. 

cohaerens (A. & S. ex Fr.) Cke.=M'. ceratopus (Pers. 1828) Qu6L 

conigenus (Pers. ex Fr.) Karst. =M. myosurus. In the Myc. myosurus comes first, followed by 
conigenus for the same species. As the latter epithet has been used by several authors for other 
species, it is better to adopt myosurus as less confusing for the brownish agaric common on 
pine cones in the autumn. 

Curreyi B. &. graminum. 

epichloe Fr. = Crinipellis stipitarius. 

erythropus (Pers.) acervatus. The epithet erythropus has also been used for other species 

and is best withdrawn. 
flosculinus (Bat.) Rea — doubtful record. 

Friesii (Bres.) Rea. =^M. j?yosurus. 
globularis Fr. = AT. WynneL 

hariolorum (DC.) QiieL=: Af. confluens. The hariolorum in the Syst. Myc. is that of BuUiard which 
is a mixture of two species. 
inodorusBat. — ^listed for Britain in error. 
ingratus (Schum.) Qu61. — appears to be M. confluens. 
msititius Rr. '=^M. VaillantiL 

von Post— doubtful record. 

(Lasch) Fr. — rnggests M. VaillantiL 
molyoidesVt. — doubtful record. 

plancus Fr. — ^not clearly defined. Cke. ///. 1073 (^9) suggests Collybia extub erans. 

polyadelpkus (JLasch) Cke. — Mycena polyadelpha. 

ruhricatus (B. & Br.) Massee — 'gTohably M. ramealis. 

saccharinus Batsch — ^identity in doubt. 

sclerotipes Bres. = Collybia cirrata vat. Cookei Bres. 

scorteus Fr, — Cke, III. lojg (1119) is form of Af. WynneL 

spodoleucus B. & Br.— probably Pteo to 

i6o Transactions British Mycologicdl Society 

MarmMim suave olem Wynnei hut iot S'poTts, 

temcellus QNulf t-x. Fr.) Karst. — in. the Sjst, Myc, both esculentus 2 ind tenacellus are used for the 
same species, but esculentus is described as growing in April and May. It is therefore the 
valid name, though tenacellus seems to be most in favour for this common spring fungus on 
pine cones. 

torquatus Fr. — doubtful record. 
torquescens Qu^L — ^also doubtful. 

undatus (Berk.) Qu^I. — ^left in, but said to be identical with M. porreus^ though the garlic smell 
is not usually detected in the fairly common species which we call M. undatus. 
mens (Bull.) Fr.=Af. permatus. Both epithets are in Syst. Myc., but mens is placed among the 
brown-spored group, most of which are Cortinarii. 
varicosus Fr. — ^has many interpretations; that of Boudier, Icon. t. 72 is usually accepted, but 
the British record is uncertain. 
xerotoides von Post — doubtful record. 

Crinipellis caulicinalis (Bull.) Rea=C. stipitarius. 



Omphalia hydrogramma (Bull.) Fr. 
umbilicata (Schaeff.) Fr. 
philonotis (Lasch) Fr. 
sphagnicola Berk, 
oniscus Fr. 

epichysium (Pers.) Fr. 
telamatiaea Berk, & Cke. 
glaucophylla (Lasch) Cke. 
leucophylla (A. & S.) Fr. 
atropuncta (Pers.) Fr. 
maura Fr. 

Pyxidatae Fr. 

Omphalia pyxidata (Bull.) Fr. 
muralis (Sow.) Fr. 
hepatica (Batsch) Fr. 
umbratilis Fr. 


var. aurea Mass, 
rustica Fr. 
chrysophylla Fr. 

Wynniae (B. & Br.) Quel. 

Alienii Maire 
striaepilea Fr. 
demissa Fr. 
rosella Lange 
Belliae Johnst. 


Omphalia umbellifera (Linn.) Fr. 
var. nivea FI Dan. 
var. flava Cke. 
var. pallida Cke. 
viridis {FI. Dan.) Lange 

Omphalia myochroa (Fr.) Rea 
grisella (Weinm.) Karst, 
retosta Fr. 
abhorrens B. & Br. 



Omphalia fibula (Bull.) Fr. 
var. Swartzii Fr. 
var. nivalis FL Dan. 
picta Fr. 

camptophylla Berk, 
griseo-pallida Desm. 

Brownii (B. & Br.) Favre. 

(See also Mycena for several species with 
decurrent gills) 

{Delicatula Fayod.) 

Omphalia integrella (Pers.) Fr. 
gibba (A. & S.) Pat. 

(iu?rom/>A<3/£;z<3 Kiihner & Maire) 

Omphalia campaneila (Batsch) Fr. 
var. papillata Fr. 
var. myriodea Kalch. 

(See also Marasmius cauticinalis) 


Omphalia gracilis Quel, 
gracillima Weinm. 
steUata Fr. 

Candida Bres. 
scyphiformis Fr. 
Mairei Gilbert 

Revised List of British Agarics and Boleti i6i 


Omphdia abhorrens 'B, 8 l Br. — not known, but its smeli and habitat is distinctive enough to justify 
its retention. 

albido-pallem — identity uncertain, and no authentic British record can be traced, 
fl/wtea Cke. & Mass. — h^xdly distinct from O, pyxidata, 

atropmcta (Pers.) Qja^l.-— left under Omphalia though Lange transfers it to Hygrophorus [CamarO'- 

Brownii (B. & Br.) Favre — transferred from Cmtharellus by Favre {Bull Soc. mycol, Fr. hV, 212) 
who cites Cke. III. 1058 (1106), so it is retained among the British records. 
buccinalis (Sow.) Cke.— a so-called common species which nobody has ever seen since Sowerby’s 

(Brig.) Cke.-— description inadequate. 
caespitosa Cke.^^O. umbelliferaYax.flava, 

campanellavdtx, badiusFT,-=^Marasmius cauiicinalis, 
detrusa Fr. — ^crept into British fungus flora by mistake, 

B. & Br.— description inadequate. 

glaucophylla {1^ Fr. — left in, but Gejp suggests it may be a terrestrial form of 0 . epichysium. 
grisea Fr.— variously interpreted but not really known. Often confused with Mycena merella 
but Fries wrote : ^ in pineti$ rara et nohilis species *, Better left out till British specimens are more 
clearly defined. 

(Batsch) Fr.— left in, but needs clearer defisnition to distinguish it from 0 . pyxidata. 
Gejp thinks them distinct. 

hydrogramma (Bull.) Fr.— this has more affinity with Cliiocyhe to which it may be transferred. 

Tho S2m.t m2iY ht of Omphalia umbilichta, 
infumata B, 81 Bt.--— not known; svLggtsi^ 0, chrysophyUd, 

Kewensis M-SiSs , — an alien on filmy fern. 

Lt^i M.a,ss,~ma.Y he 3. form of Clitocybe Jragrans* 

Berk.— an alien in orchid house. 
offuciata Br.-^not known; almost suggests Laccaria laccata, 
pseudo-androsacea (Bull.) Fr, = 0 , umbellifera, 
pseudo-directa W.G.Sm. — ^no spores given. 
scyphoides Fr.—-douhtfnl identity; prohahly Clitapilus cretaius. 
spkagnicola Berk.— this rare species is (sec. Gejp) distinct from 0 . philonotis. 
telmatiae Berk. & Gke.— left in list for the present, but may be large form of 0 . philonotis* 
tricolor (A. & S.) Fr.^Marasmius tricolor, 
umbelUfera \2ir. citrina Qutl.^ 0 . Wynniae, 
delutina QuB,^ 0 . grisella. 


Pleurotus ostreatus (Jacq.) Fr. 
var. columbinus (Quel.) Gke. 
var. salignus (Pers.) Fr. 
var. etiosmus (Berk.) Gke. 

^ cornucopiae (Paulet) Persoon 
ulmarius (Bull.) Fr. 
porrigens (Pers.) Fr. 

Hgnatilis Fr. 
dryinus (Pers.) Fr, 
mutilus Fr. 

{Acanthocystis Fayod) 

{a) Cystidia present 

Pleurotus petaloides (Bull.) Fr. 
var. geogenius (DC) Pilat 
mastrucatus Fr. 
reniformis Fr. 

Pleurotus atrocaeruIeus Fr. 
serotinus (Schrad.) Fr. 
var. Almenii (Fr.) Big, & GuilL 

{b) Cystidia absent 

Pleurotus applicatus (Batsch) Fr. 
Silvanus Sacc. 
cyphelliformis Berk. 

Lauro-cerasi B. & Br, 

Leightonii Berk. 

{Pleurotellus Fayod) 

Pleurotus chioneus (Pers.) Fr. 
septicus Fr. 

hypnophilus Berk, sensu Sacc., Rea, 
etc., non Quel, 
dictyorrhizus (DC) Fr. 



Transactions British Mycological Society 

(Pkurotellus Fayod) {continued) 
Fleurotus acerosus Fr. 
tremulus (SchaefF.) Fr, 
candidissimus Berk. & Curt. 

Pleurotus mitis (Pers.) Berk. 

{Rhodotus Maire) 
Pleurotus palmatus (Bull.) Fr. 


Pleurotus acerinus'Fv.^P^ dryinus. 
acerosus stmu.'R.tdi=^P, tremulus, 
algidusPt.^P, atrocaeruleus {stc. Pilot). 

applwatus — ^with elliptical spores sensu Rea is P, Silvanus which we have listed. P. applicatus 
has globose spores and as it is said to be a common species, we have not removed it. 
circinatus Fr. — ^probably P. lignatilis, 
corticatusPx.=P. dryinus. 

craspedius Fr.— doubtful species. Gke. III. 274 (256) is P. ulmarius. 
fimbriatus {"Rolt.) Pi. ^ ioxmo^ P. lignatilis. 

Jlicxzlis Fr. — doubtful; Pilat suggests it may be P. unguicularis Fr. 

Hobsonii Berk. — probably P. chioneus. 

hypnophilus Berk. — this epithet has been applied to two species. It is left in the list as applying 
to the species with minute spores. Pilat adopts the interpretation of Quelet, the spores given 
being 6-8 x 2*6-3//-, may be a form of P. septicus. 

Leightonii Berk. — also left in list, but Cke. III. 290 (260) is P. atrocaeruleus. 
limpidus Fr. — doubtful species. 

pantoleucus Fr.=P. dryinus. Cooke’s figure 277 (179) is either Panus torulosus or Pleurotus paU 

pulmonariusPt.=P. ostreatus. 
revolutus Pilch:.. = P. ostreatus. 

rufipes Mass. & Smith— Pilat says this may be P. mitis with stem coloured by contact. 

Ruthae B. & Br.— form of P. petaloides according to Pilat who examined specimens at Kew. 

sapidus Sch.\x\zcx 3 iQ']^=P. comucopiae {Petvlct) Pcxsoon, 1^2^. 

spongiosusPx.^P. dryinus. 

striatulus Fr.=P. applicatus. 

tessulatus (Bull.) Fr.=P. ulmarius, 

tremulus scnsvi'R.tdi—P. acerosus. 

Panus torulosus (Pers.) Fr. 
rudis Fr. 

Gen. PANUS Fr. 

{Panellus Karst.) 
Panus stipticus Karst. 

var. farinaceus (Schum.) Rea 


Lentinus tigrinus (Bull.) Fr. 
lepideus Fr. 

adhaerens (A. & S.) Fr. 

{Lentinellus Karst, em. Kuhner) 
Lentinus cochleatus (Pers.) Fr. 

Lentinus omphalodes Fr. 

var. scoticus (B. & Br.) Pilat 
vulpinus (Sow.) Fr. 
f. auricula (Fr.) Pilat 


Schizophyllum commune Fr. 

Nyctalis parasitica (Bull.) Fr. 


Nyctalis asterophora Fr, 

Revised List of British Agarics and Boleti 163 


Panm comhattis (Bvlh) torulosus. 

paUllaris Fr.— British record doubtful. Gke. Ill, 1097 (1144) has spores 4x3/4, but Pilafs 
measurements are 3-4 X o-8-r *3 /4. 

StevensoniiB, &L'Bt,=^Pleurottis nidulam, 

Xerotus degenet Yr.—Lentinus cyathiformts^ but Cke. Ill, 1098 (1150) looks like Collybia clusilis, 
Lentinus auricula Pt, — fotta of L, vulpinus, 
fimhriatus Gurrey=X. tigrinus, 

flabellinus Q)x€l,=L, omphalodes, 

leontopodius Schulz. cyathiformis, but the British record of this rare species is doubtful. 
pulverulentus {Scop.) PT.=iL. adhaerens, 
scoticus B. & Br.=var. of L, omphalodes, 
sulcatus Berk. 

suffrutescens (Brot.) Fr.— probably L, lepideus, 
umbellatusPt.=L, cochleatus. 




1 , {LimaciumPx,) 

A. Candidi Bat. 

Hygrophorus eburneus (Bull.) Fr. 
cossus (Sow.) Fr. 
chrysodon Fr. 
penarius Fr. 
discoxanthus (Fr.) Rea 

B. Pudorini Bat. 

Hygrophorus Russula (Schaeff.) Quel, 
erubescens Fr. 
pudorinus Fr. 

C. Luteoli 

Hygrophorus lucorum Kalch. 
discoideus (Pers.) Fr. 
leucophaeus (Scop.) Fr. 
arbustivus Fr. 
aureus (Arrh.) Fr. 

D. Olivaceo-umbrini Bat. 

Hygrophorus olivaceo-albus Fr. 
var. obesus (Bres.) Maire 
limacinus (Scop.) Fr. 
squamulosus Rea 
fusco-albus Fr. 
cerasinus Berk, 
hypothejus Fr. 

E. Griseoli 

Hygrophorus agathosmus Fr. 
var. aureo-floccosus Bres. 
pustulatus (Pers.) Fr. 
livido-albus Fr. 

II. (Camarophyllus Fr.) 


Hygrophorus camardphyllus (A. & S.) Fr. 
nemoreus (Lasch) Fr. 
pratensis (Pers.) Fr. 

var. pallidus B. & Br. 
cinereus (Fr.) Karst. 

Karstenii Sacc. & Cub. 
virgineus (Wulf.) Fr. 

var. roseipes Mass, 
niveus (Scop.) Fr. 
russo-coriaceus Berk, & Miller 
subradiatus (Schum.) Fr. 
lacmus Fr. 

Golemannianus BIox. 


Hygrophorus fornicatus Fr. 
citrino-virens Lange 
ovinus (Bull.) Fr. 
metapodius Fr. 
lepidopus Rea 

III. {Hygrocybe Fr.) 

{a) Conici'BB.t, 

Hygrophorus conicus Fr. 
nigrescens Qjuei. 
intermedins Pass, 
obrusseus Fr. 
calyptraeformis Berk, 
var, niveus Gke. 

1 1-2 

Transactions British Mycological Society 


{b) Coccinei Bat. 

Hygrophoms coccineus (Schaeff.) Fr. 
puniceus Fr. 
miuiatus Fr. 

Real Maire 
mocronelius Fr. 
sdopbanus Fr. 
sciophanoides Rea 
tunindus Fr. ‘ 
var. mollis B. & Br. 

(c) Flaveoli 

Hygrophorus psittacinus (Schaeff.) Fr. 
ceraceus (Wulf.) Fr. 
citrinus Rea 
chlorophanus Fr. 
vitellmusFr. ; 
laetus (Pers.) Fr. 
micaceus B. & Br. 

{d) Tristes Bat. 

Hygrophorus unguinosus Fr. 
nitratus (Pers.) Fr. 
foetens Phill. 


Hygrophorus amoenus Lasch (1828)— most continental authors synonymize this with H. calyptrae- 
formis Berk, (i860), but the original description by Lasch in Linnaea^ voL iii, p. 390, gives 
the essential characters as ^PiUo. . ,ohtusOy struito^ suhaurantiacay lamellis decurrentibus^ y all of 
which are different from H, calyptraeformis. 
aromatkm (Sow.) Berk. — ^may be a form of H. laetus. 

Clarkii B. & Br.=if. unguinosus. 
connatusK.-aist.-^’^ form oi H. unguinostts. 

glutinifer Fr. — variously interpreted. Cke. III. 878 (889) is probably H. olivaceo-albus. 

irrigatus unguinosus. 

leporinus Fr. = H. nemoreus. 

mlizeus’Ft . — does not seem distinct from if. 

mesotephrus B. & Br.=i/. leucophaeus. 

niveus {Scop.) Vr. vsLT.fuscescens subradiaius. 

obscuratus Karst.— probably H. nitratus. 

persicinus Beck. — suggests a. form of Cantharellus dbarius. 

pratensis var. cinereus Fr.=if. cinereus. 

pulverulentus B. & Br. — ^known only from three rather dubious old records. 
sciophanoides Bea : — ^left in, but may be if. 

spadiceus (Scop.) Fr.— record doubtful. Cke. III. 1194 (1161) with its small spores is probably 
H. hypothejus. 

squamuhsus Rea — ^left in, but suggests a Tricholoma near sejunctum. It was originally recorded 
from Clare Island, west of Ireland. 
tristis (^ers.)Bres.^H.nigrescens. 

turundus var. sphaerosporus Rea— this can hardly be correct. The spores as figured in original 
drawing are those of a Laccaria. 
ventricosus B, %L Br.— form of H. virgineus. 

WynniaeB. & Br. = Omphalia Wynniae. 


I. Glutinosi Qudl. 

A. Barbati Quel. 

Lactarius scrobiculatus (Scop.) Fr. 
resimus Fr. 
repraesentaneus Britz. 

Lactarius torminosus Fr. 
cilicioides Fr. 
pubescens Fr. 

plumbeus Fr. ( = turpis Fr. ) 
controversus (Pers.) Fr. 

Revised List of British Agarics and Boleti 

B. Glahrati Batailie 

(i) Color enti 

Lactarius deliciosus (Linn.) Fr, 
uvidus Fr. 
vioiascens Fr. 
aspideus Fr. 
chrysorheus Fr. 
vietus Fr. 
blennius Fr. 
umbrinus (Pers.) Fr. 

(See also L, theiogalus) 

(2) Immutabiles 

Lactarius zonarius (Bull.) Fr. 
insuisus Fr. 
pyrogalus (Bull.) Fr. 
fiexuosus Fr. 

var. roseo-zonatus Fr. 
circeilatus (Batt.) Fr. 
hysginus Fr. 
triviaiis Fr. 
pallidus (Pers.) Fr. 

11 . Velutini Opel, 
(i) Albati^aX, 

Lactarius vellereus Fr. 
piperatus (Scop.) Fr. 

(2) Rubescenti 
Lactarius fuliginosus Fr. 
picinus Fr, 

lignyotus (Lindb.) Fr. 
acris (Bolt.) Fr. 

(3) Olenfes 
Lactarius helvus Fr. 
camphoratus (Bull.) Fr. 
serifluus (DC.) Fr, 
cimicarius (Batsch) Cke. 
quietus Fr. 
giycyosmus Fr. 

(4) Acri 

Lactarius rufus (Scop.) Fr. 
mammosus Fr. 
iilacinus (Lasch) Fr. 

(5) Subdulces Konr. 
Lactarius volemus Fr. 
ichoratus (Batsch) Fr. 
subdulcis (Pers.) Fr. 
aurantiacus {FL Dan.) Fr. 
mitissimus Fr. 
cremor Fr. 

cyathuia (Fr.) Ricken 
obnubilus (Lasch) Fr. 
theiogalus Fr. non Bull, 
hepaticus Plow. 



Lactarius (Bolt.) Fr. — ^left in with some doubt, as it hardly differs from L, fuliginosus. 
capsicum Schulz. — ^not known. 

chrysorheus Fr. Epicr. 1836--8. There can be little doubt that this is the same as £. theiogalus 
(Bull.) Fr. Syst. Myc., though in later works Fries identified the latter name with a zoneless 
agaric, the milk of which tardily turns yellow. To avoid confusion we have left both epithets 
in the list, 

circeilatus "Bt. — ^it is not clear whether this differs from 
flavidus Bond, aspideus. 

fiexuosus Bv. — nolo on L, circeilatus. Both kept in list. 
fiuens Bond. blennius. 

Soppitt. — spores in Gke. ///. 1 195 (1194) not of a 
lateritiorosem Bi'axst.^^iL. Iilacinus. 

limdus Lamb.~not known and description inadequate. 
mammosus Br. — -left in, but hardly known. 
minimus W.G.Sm. — dwsid form of L. subdulcis. 
obliquus Br.— -not known. Gke. ///. 969 (1014) may be form of L. 

obnubilis (Lasch) Fr. — ^left in but variously interpreted, so a clear description of British record 
is needed. 

pergamenus Swartz^ L. piperatus. 

representaneus Britz. — ^very like L. scrobiculatus but milk turns violet. Found at Rothiemurchus, 
Scotland, some years ago, when it was listed as £. aspideus. 
resimus Fr. — doubtful if distinct from L. scrobiculatus, but left in for further study, 
retisporus Mass.— dark form of L. fuliginosus. 
sanguifluus (Bsiul.) Br. — a southern European species. 

i66 Transactions British Mycological Society 

* Lactarius scotkus B. & Br. — figure in Ckc. IlL 938 (1004) suggests L, pubescens, 
spmomius=L. lilacinus* 

squalidns (Krombh.) Fr. — of dubious identity. 

subumbomtus Lindgr. — considered to be either L. cimicarius or Z. camphoratus. 
tomButosus {Otto) CkQ, — not yrdb^bly L, torminosus, 

tabidus Fr. — variously interpreted. Sensu Boudier=X. cyathula, 

turpis FT.=plumbei£S Fr. Myc. (i8f2i). Much as we dislike disturbing this familiar name 
plumbeus has priority. Another name for the same species, necaior Pers., also appears in the 
Syst. but has been confused with torminosus. It is therefore best to adopt plumbeus, though 
Bresadoia discarded it because it is a ^ nomen ineptum'; but that is not a valid reason for 
excluding it. 

umbrinus (Pers.) Fr. — ^remains in list, but this rare and little known species has not been found 
for years. 

utilis (Weinm.) Fr. — doubtful. Cke. IlL 930 (1084) may be L, delica. 


I. Lactariodes {b) Pallidisporae 

Russula nigricans Fr. 
densifolia (Seer.) Gill 
adusta Fr. 
albo-nigra Krombh. 
delica Fr. 

11 . Rigidae Fr. 

Russula mustelina Fr. 
virescens (Schaeff.) Fr. 
lepida Fr. 

var. amara Make 
lactea (Pers.) Fr. 
rosea Qu 61 . 

f. aurora Pearson 
azurea Bres. 

Russula violacea Quel, 
sanguinea (Bull.) Fr. 
Queletii Fr. 
dreimeia Cke. 

var. viridis Singer 
fellea Fr. 

{c) Xanthosporae 

Russula rubra (Kromb.) Bres. 
badia Quel, 
maculata Quel, 
luteo-viridans Martin 
nitida (Pers.) Fr. 

amoena Quel. 

III. Resilientes 

Russula cyanoxantha (SchaefF.) Fr. 
heterophylla Fr. 

var. virginea (Cke. & Mass.) comb. nov. 
vesca Fr. 
farinipes Romell 

VI. Sub-acrae 

Russula atro-purpurea Kromb. 
var. depallens (Cke.) Make 
ochroleuca Fr. 

Solaris Ferd. & Winge 
pseudo-integra Arn. & Goris 
veternosa Fr. 

IV. Foetentes Kiihner & Joss. 
Russula foetens Fr. 
iaurocerasi Melz. 

exalbicans Bres. 
versicolor J. Schaeffer 
gracillima J. Schaeffer 

pectinata (Bull.) Fr. 
sororia Fr. 
consobrina Fr. 

V. Acrae 
(a) Lemosporae 

Russula emetica (Schaeff.) Fr. 
Mairei Singer 
luteo-tacta Rea 
fragilis Fr. 

var. nivea (Pers.) Cke. 
fallax (Fr.) Cke. 

VII. Gratae 
{a) Pallidisporae 

Russula grisea (Pers. ex Seer.) Fr. 
aemginea Lindb. 
melliolens Quel. 

var. Chrisma.ntiae Maire 
puellaris Fr. 

var. leprosa Bres. 
brunneo-violacea Crawshay 
lilacea Qudl. 

Revised List of British Agarics and Boleti 

(b) Xanthosporae 
Russula xerampeiina Fr. 

var. fusca (Qiiel.) Melz. & Zv. 
var. erythropus Pelt, 
var. olivascens (Fr.) Melz. & Zv. 
decoiorans Fr. 
claroflava Grove 
vinosa Lindb. 
alutacea (Pers.) Fr. 
olivacea (SchaefF.) Fr. 

Romeliii Maire 
Integra (Linn.) Fr. 


Russula caeruiea Gke. 

Turci Bres. 

Velenovskyi Melz. & Zv. 
paiudosa Britz. 
aurata (With.) Fr. 
iutea Fr. 

f. iuteorosella Britz. 
venosa Vel. 

Scotica Pearson 
nauseosa (Pers.) Fr. 

(See wid.tT Sub-acrae for species that may 
be mild or acrid) 


Russula armeniaca Cke. =i 2 . Iutea var. Iuteorosella Britz. 
aurora Krombh. — there is some doubt whether this is a variety of R, lepida or identical with 
R, rosea Quel. It is left in the list as a peach-coloured form of R. rosea^ as described in Trans. 
BriL mycoL Soc. xxn, 36. 
carnicolor Brts.^R. lilacea. 

chamaeleontinaBx . — variously interpreted; not clear what the British records refer to. Gke. III. 

1054 (1098) is a mixed bag. 
citrina GtIIgX — bright yellow form of i2. 
eonstaas (Karst.) Romell=i 2 . clarqflava. 

cutifracta Oh &. — ^a much disputed species; nobody knows what it really is. 
drimeia Cke. — ^many authors adopt R. sardonia Fr. for this species, but as Fries states that the 
gills are white, we prefer to retain Cooke’s epithet for the species with primrose gills. 
drimeia var. Jlavo-virens Rea (1932) =R. drimeia f. viridis Singer (1928). 
elegans Bres.-~? form of R. xerampeiina. 
elephantina'Fv.‘=R. delica. 

emetica Fr.— -there are conflicting opinions about this species. The beechwood form is recorded 
in this list as R. Mairei Singer, but the other forms require further study before they can be 
definitely placed. 

fingibilis Britz. =R. ochroleuca^ which may be mild or acrid. 

fragilis (Pers.) Fr.— there is no universal agreement about this species. It is often identified 
with a scarlet-capped Russula of the emetica group, but this can hardly be correct. Persoon 
described it as pileo rubro purpurascente. Fries in Syst. Myc. as e purpureo exalbicat. In Hym. Eur. 
Fries states that it often has an eroded edge to the gills. It seems, therefore, reasonable to 
identify the species with the small fragile Russula with a purplish red pileus and fimbriate 
gills which is so common in Britain. 
furcata (Pers.) Fr. — usually considered as a green form of cyanoxantha. 
fusca Qu^I. = either i2. mustelina or a form of R. xerampeiina. 
gahchroaEr.^R. heterophylla. 
graminkolor {f>oor.) aeruginea. 

incarnata (^€i.-=R, rosea 

Fr.— a large form of i 2 . 
mitis 'Rj&dL= R. vesca. 

ochracea (A. & S.) Fr.— variously interpreted; usually considered to be R.fellea, 

(Gillet) Maire Qud. 

rosacea (Bors.) Er.=R. sanguinea. 

roseipes (Seer.) Bres,— doubtful British record. It is quite distinct from R, roseipes Gke. III. 1035 
( 1 08 1 ), which is R. (Fr.) J. Schaeffer. 

semi-crema Bv.^R. densifolia. 
serotina Qn €\. — ^it is not clear what this is. 

sphagnophila Kauffm.— this epithet was applied by Singer to a common and very variable 
Russula which grows in wet places usually imder birch trees. Many, names have been given 
to this species, and it was thought that J?. venosa fyd.) Melzer would be generally accepted. 
J. Schaeffer in Ann. Myc. xxxviii (1940) identifies it with the true R. mtida Fr. For the 

l68 Transactions British Mycological Society 

present we prefer to leave R, verwsa as the epithet which clearly identifies this common 
Russula, Sind R, nitida remains to indicate the acrid species which hitherto has been associated 
■ with , this name, ■, 

Russula subfietensW. G. Sm . — probably a form of R.foetens, but it is not known what it really 
refers to. R. subfoetens sensu Maire and also Melzer is R. farinipes Romell. Gke. IlL ioi6 
f 1 047) looks like 

mrgma Ckt. 8c M.2 lBs, = ybx. of R. heterophylla, 

violacea Qu6L — though marked as ‘common’ by Rea the authentic R, violacea with its smell of 
laudanum is rare in Britain, and the agaric often determined under this name is either R. 
or another species with a violet pileus. 


Gantharellus cibarius Fr. 
var. aibus Fr. 

var. amethysteus (Quel.) Maire 
Friesii Quel. 

carbonarius (A. & S.) Fr. 

Gantharellus tubaeformis (Bull.) Fr. 
lutescens (Pers.) Fr. 
cinereus (Pers.) Fr. 
cupuiatus Fr. 


Neurophyllum clavatum (Pers.) Pat. 

Gen. LEPTOTUS Karst, em. Maire 

Leptotus muscigenus (Bull.) Maire Leptotus lobatus (Pers.) Karst, 

retirugis (Bull.) Karst. glaucus (Batsch) Maire 


Graterellus cornucopiodes (Linn.) Fr. Graterellus amethysteus Rea 

sinuosus Fr. 


Plicatura crispa (Pers. ex Fr.) Rea 


Canikarellus albidus Fr. = Clitocybe albida, 
aurantiactis (y^xslt) 'Fx. = Clitocybe aurantiaca, 

BromniiB. 8c '&v,^0mphalia Brownii. 

HoughtoniiBhiVi . — Gke. IlL 1060 (1107) looks like a form of Mycena galericulata* 
infimdibulifomis (Scop.) Fr. = C, iubaejwmis. ' 
l^ophaeus Nouel. = C. carbonarius. 
replexus Fr.— doubtful identity. 

StevensoniiB. 8c Bx.=- Clitocybe albida. 
umbomtus (Gmel.) Fr. = Clitocybe umbonata. 

Trogia crispa Plicatura crispa. 

Quel. (1886)= Karst. (1879). 

Graterellus crispus Pt.^C. sinuosus. 

pusillus Fr. = G. sinuosus. 


A. Viscidae B. Sericellae 

Volvaria speciosa Fr. 

f. gloiocephala (DG.) Konrad & 
media (Schum.) Fr. 
media biloba Mass. 

Volvaria pusilia (Pers.) Fr. 
voivacea (Bull.) Fr. 
Taylori Berk, 
bombycina (Pers.) Fr. 
surrecta (Knapp) Ramsb. 
murinella Quel. 


Revised List of British Agarics and Boleti 


I, Tricholodermei Lange 
A. Coronati Lange 

Pluteus cervinus (SchaeiF.) Fr. 
var. atromarginatus Konr. 
var. patricius (Schulz.) Fr. 
var. eximius (Saund. & Sm.) comb. nov. 
Salicinus (Pers.) Fr. 
petasatus (Pers.) Fr. 
pellitus (Pers.) Fr. 

B. Depauperati Lange 

Pluteus Buiiii Berk, 
gracilis (Bres.) Lange 
hispiduius (Fr.) Quel. 

Pluteus umbrosus (Pers.) Fr. sensu Ricken 
iuteo-virens Rea 

II. Micacei Lange ■ 

Pluteus semi-bulbosus (Lasch) Fr. 

Godeyi Giilet 
cinereo-fuscus Lange 
nanus (Pers.) Fr. 

var. iutescens Fr. 
chrysophaeus (SchaefF.) Fr, 
phlebophorus (Ditmar) Fr. 
marginatus (Quel.) Bres. 
argenteo-griseus Rea 
leoninus {SchaefF.) Fr. 
coccineus (Mass.) Lange 
vioiarius Mass. 


Pluteolus aleuriatus Fr. 

var. reticulatus (Pers. ex Fr.) Lange 


Fa/zjflm Berk. =: F. (Knapp) Ramsb- 

media (Schum.) Fr. — ^is retained, but the identity of the British record is not clear. Lange 
gives the spores as I i-i 6 X 7-8 jtt. Rea has 5-6x4 /u. 
murinella Qu^L— recorded hi j, Bot, lxx (1932), but without microscopic details. 
parvula (Weinm.) ¥r. ~V. pusilla, 
tmperata B. & Br. — hardly distinct from F. pusilla* 
viperina Fr. — -British record doubtful. 

Huteus cervinus — ^the usual varieties are left in, but they are mostly forms that occur on sawdust. 
ephebius Fr.^ — ^not clearly defined. Gke. Ill, (517) looks like a rather large Pluteolus aleuriatus, 
melanodon (SecrJ) Fr. — doubtful identity. 
nanus yax, major Mass.=P. cinereo-fuscus. 
roseo^albzis Ft. — ■inadequately described. 
spilopus B, & Br. — ^may be a form of P. cervinus. 
sororiatus — ^not known; may be a 

(Sommerf.) Fr. — ^appears to be a pale form of P. nanus, 
vioiarius Mass. — ^left in because it has distinctive characters, but it has not been found since 
first collected. 

Pluteolm Mulgravensis yia$s. 81 Cxo^s\.---iptohBb\Y P- aleuriatus. 


Clitopilus prunulus (Scop.) Fr. Glitopilus cretatus B. & Br. 

Gen. RIPARUTES Karst. 
Ripartites tricholoma (A. & S.) Karst. 

lyo Transactions British Mycological Society 


ClUopUm ¥r,--~thm genus is now confined to species with ribbed or polygonal pinkish spores^ 
angustus Fr.— not known what this refers to. 

carneo-albns ‘Bv.^^dQcntvtntiovmoi Leptonia sericella. 


mmdulus (Jjsisch) 'Br.^^Clitocybe {Rhodopaxillus) mundula. 

orcdla (Bull.) Fr, = C, prmulns. 

popinalis 'Ft. = Clitocybe {Rhodopaxillus) popinalis. 

Sarnicus Mass, — probably decurrent form of Entoloma ardosiacum=^ Eccilia Mougeotii. 

Smithii Mass. — Gke. III. 350 (599) looks like Laccaria laccata. 

stilbocephalus B. & Br. — ^identity uncertain; seems to be an Entoloma. 

straminipes M. 3 .?,$.^Eccilia straminipes. 

undatus Ft. — Eccilia undata. 

vilis Ft. — Eccilia undata or E. Mougeotii, 

[Rhodophyllus QnS..) 

1. Gen. ENTOLOMA Fr. 


Entoloma porphyrophaeum Fr. 
jubatum Fr. 
dichroum (Pers.) Fr. 
excentricum Bres. 
erophilum Fr. 
griseo-cyaneum Fr. 

var, roseum Maire 
turbidum Fr. 

Entoloma lividum (Bull.) Fr. 
prunuloides Fr. 

Entoloma nitidum Quel, 
ardosiacum (Bull.) Fr, 
var. Mougeotii (Quel.) 
madidum Fr. 

Bioxami B. & Br. 
clypeatum (Linn.) Fr. 
sericeum (Bull.) Fr. 
costatum Fr. 
rhodopolium Fr. 
nidorosum Fr. 
ameides B. & Br. 
speculum (Fr.) Quel, 
nigrocinnamomeum Kalcli. 

Leptonia lampropus Fr. 
placida Fr. 
anatina (Lasch) Fr. 
lappula Fr. 

Reaae Maire 
aethiops Fr. 
solstitialis Fr. 
sericella (Fr.) Quel. 

var. decurrens Bond, 
incana Fr. 

Formosa Fr. 


Leptonia Formosa 

var. suavis (Lasch) Fr. 

Queletii Boud. 
chalybaea (Pers.) Fr. 
serrulata Fr. 

var. laevipes Maire 
var. atrides (Lasch) Kiihner & Maire 
lazulina Fr. 
euchroa (Pers.) Fr. 
asprella Fr. 
nefrens Fr. 
sarcita Fr, 


Nolanea staurospora Bres. 
cetrata (Fr.) Schroet. 
hirtipes (Schum.) Fr. 
mammosa (Linn.) Fr. 
papillata Bres. 
infula Fr. 
juncea Fr. 
versatilis Fr. 
minuta Karst. 

Nolanea icterina Fr. 
Babingtonii Blox. 
fulvo-strigosa B. & Br. 
fumosella Winter 
coelestina Fr. 
exilis Fr. 
verecunda Fr. 
rufo-carnea Berk, 
vinacea (Scop.) Fr. 

Revised List of British Agarics and Boleti 171 


Eccilia cancrina Fr. Eccilia parkensis Fr. 

carneo-grisea B. & Br. straminipes (Mass.) comb. nov. 

griseo-rubelia (Lasch) Fr. decurrent forms oi Leptonia 

rhodocylix (Lasch) Fr. Entoloma) 

undata (Fr.) Big. & Guiii. 

V. Gen. CLAUDOPUS W.G.Sm. em. Patouiilard 

Giaudopus depluens (Batsch) Fr. Claudopus byssisedus (Pers.) Fr. 

(see Crepidotus foT CL variabilis, etc.) 


Rhodophyllus Quel. — ^some modern authors have adopted this genus to include all the agarics 
with pink nodulose spores, but the old Friesian sections are still serviceable. Romagnesi 
has worked out a new grouping largely based on the shape of the spore, which is very im- 
pressive but too complex for general use. 

Entoloma Batschianum Fr. — doubtful British record. 
bulbigenum B. Si Br, — prohsihly Leptonia sericella. 

Farrahii . — ^has smooth spores, so not an Entoloma, 
fertile Berk, ^ E, lividum. 

helodes Fr. — ^not known. Cke. Ill, 322 (329) suggests E. prmuloides, 

jubatum Fr. — often confused with E, porphyrophaeum which is very common. E. jubatum is 
smaller and has no trace of purple. 
liquescens Ckc.— Psilocybe sarcocephala or P. spadiceo-grisea, 

majale Fr. — doubtful British record. Nolanea majalis (Fr.) Konr. was renamed jV. hirtipes. 
nigrocinnamomeum Kalch, — ^left in because Massee^s figure in Cke. III. 1158 (1153) is a well- 
marked agaric difficult to place elsewhere. 

placenta (Batsch) Er.—E, porphyrophaeum sec. Konr. & Maubl. but Cke. III. 321 (314) looks like 
a small Collybia platyphylla. 

pluteoides Fr. — ^identity uncertain; may be large Pluteus semi-bulbosus. 

pulvereum Nolanea fumosella, 

repandum Fr. — ^hardly distinct from E, prunuloides, 

resutum Fr. — doubtful record. 

Rozei Qu 61 . — doubtful record. 

Saundersii~- 4 rom W. G. Smith’s figure in Cke. III. 331 (306) and the spores given by Rea, this 
is the white form of Pluteus cervinus [patricius) , Boudier has spores i i-i 3 ft, but his is a different 


Thomsonii B. Si Br. — position uncertain — doubtful if an Entoloma or a Pluteus. 
torfipes Mass. — the description and spores suggest typical Collybia distorta, 
venosum Gillet — probably E. costatum. 

Wynnei B. & Br. — may be E. griseo-cyaneum. 

Eccilia acus VI.G.Sra , — an alien on coffee seeds in greenhouse. 
atro^puncta Er.^Omphalia atro-puncta. 

flosculus .G.Bm . — an alien in fern house. 

nigrella (Pers.) Gillet — deairvont form oi Leptonia serrulata. 
j&iirtem Fr.—left in, but not recorded in Britain for many years. 

Leptonia acuta Kosi — ^.n Sicute form of Entoloma sericeum. 
chloropolia Fr, — -prohsihly JVolanea icterina. 
euchlora (Lasch) Fr. =X. fwfl«tz. 
serrulata var. Berkeley Entoloma ardosiacum. 

Nolanea araneosa Q,ueL=JV- fulvo-strigosa B. & Br. sec. Bresadola. 
majalis (fr.) l^orir,=='M. hirtipes. 
nigripes (Trog) Ft, ==- Maucoria cucumis. 

pascua (Pers.) Fr.— a collective species. It was described by Fries in Syst, Myc. as proteus hujus 
sectionis quern in decern species distinguere postis. In Britain it was until recent years applied to 
a very common which has been clearly defined by Bresadola tmder the name 

and there seems to be general agreement to accept this. 

172 Transactions British Mycological Society 

Molaneapiscmdora (Ces.) Fr.— usually considered as the same as Naucoria cucumis, though the epithet 

has been applied to a genuine 
proktaria ^ovd,=^N. staumpom, 

rhodosptyra Br, & W.G.Sm. — an alien. 

rubida Berk.— not a Nolanea. Cke. Ill 367 (340) looks like CUtopilus cretatus. 
rufo-mmea Berk.— not known. Cke. Ill 364 (378) is copied from Berkeley’s original drawing 
but so badly reproduced that the plate looks like Laccaria laccata, quite unlike the original 
painting which represents an unfamiliar agaric. 

Rea Winter. 

subglobosa A. & S.— Cke. Ill 1160 (1170) may be jV*. icterina though the smooth spores suggest 
a Pluteus. 

Claudopus varkhilis and sphaerosporus — the genus Claudopus is applied by most modern authors 
only to species with pinkish angular spores. Cl variabilis has smooth spores which are clay- 
coloured in mass, and it is better to restore it to its original position as a Crepidotus together 
with its roimd-spored ally. 



Pholiota squarrosa (Mull.) Fr. 
subsquarrosa (Mull.) Fr. 
Mulieri Fr. 
aurivella (Batsch) Fr. 
grandis Rea 
adiposa Fr. 
lucifera (Lasch) Fr. 
spectabilis Fr. 
flaxnmans Fr. 
mutabilis (SchaefF.) Fr. 
terrigena Fr. 
curvipes Fr. 

tuberculosa (Schaeff.) Fr. 
destruens Brond. 

{Phaeolepiota Maire) 
Pholiota aurea (Mattusch.) Fr. 

{Rozites Karst.) 
Pholiota caperata (Pers.) Fr. 

{Hebeloma Fr.) 

Pholiota radicosa (Bull.) Fr. 

{Naucoria Fr. Phaeomarasmius ScherfF.) 
Pholiota erinacea (Fr.) Quel. 

V {Agrocyhe'E2iyodi, Togaria 
W. G. Sm. em. Romagnesi) 
Pholiota praecox (Pers.) Fr. 
dura (Bolt.). Fr. 
sphaleromorpha (Bull.) Fr. 
aegerita Brig, 
erebia Fr. 

[Conocybe Fayod em. Kiihner) 
Pholiota togularis (Bull.) Fr. non Ricken 
blattaria Fr. non Ricken 

{Galerina Kiihner) 

Pholiota marginata (Batsch) Fr. 
mycenoides Fr. 
pumila Fr. 


Pholiota confragosa Fr. — a little-known species; British records not certain. 
cruentatus Cke. & Sm. — the figures in Cke. ///. 399 (366) may be Cortimrius [Inoloma) rubicun- 
(Rea) Pearson. 

dissimulans B* Bn ^T.=^sm2i\\ P, erebia, 

grandis Rea— said to be not uncommon but not well known. Left in, but doubtful whether 
different from a large P. 
heteroclita Br, — Cke. Ill $Sg {^6S) —P, destruens. 

Junonia Fr. — Cke. Ill 397 (369) represents a slender form of P. spectabilis. 
leochroma Cke. =P. aegerita. 
molliscorium Cke. Mass. =P. erebia. 

mwrkata Fr. — this species has been referred to Naucoria, but we do not know if British record 
refers to the same agaric. 

mustelina Fr. — position uncertain so left out, but Cke. Ill 404 (356) looks authentic. 

ombrophila Br.=P. erebia. 

paxillusBr. — ^British record doubtful, ^ ^ ^ ^ ^ 

Revised List of British Agarics and Boleti 173 

Pkoiiota phalerata Fr,— -probably form of F. caperata. In , any case the British record was due to 
a mistake. 

rujidula Kalch.^ — may be Tubarmfurfuracea; no authentic British collection. 
sororia Karst. — ^requires clearer diagnosis. 
sublutea (FL Dan,) Ft . — ^not adequately described. 
micolor {FL Dan.) Fr. — ^hardly distinguished from P, marginata, 

Alexandri Gilltt — ^recorded as British in error. 
acetahidosa {Sow.) Sacc.-— doubtful species. 


Demdata Fr. 

Hebeloma sinuosum Fr. 
sinapizans (Paul.) Fr. 
crustuliniforme (Bull.) Fr. 
iongicaudum (Pers.) Fr. 
elatum (Batsch) Fr. 
anthracophilum Maire 
sacchariolens Quel, 
subsaponaceum Karst. 


Hebeloma capniocephalum (Bull.) Fr. — ^uncertain species; Cke. III. 462 (419) looks like H. meso- 
phaeum. Bresadola interprets Bulliard’s species as a Collybia but his figures on PI. 199, Icon. 
Myc.y bear little relation to those of Bulliard. 
claviceps Fr. — probably a form of H. versipelle. 
dif ractum Ft. ^H. crustuliniforme. 
firmum Fr. — hardly distinct from H. testaceum. 
glutinosum (JAndgt.) Ft.^Flammula lenta. 
ichnostylum Cke. = /f. sacchariolens. 
lugetis (Jungh.) Fr, — ^not known. 
magnimamma Fr. — doubtful to what this refers. 

mussivum Fr. — probably Cortinarius percomis. The figures in Fr. Icon, m suggest this. 
nauseosum Cke, — may be H. sacchariolens. The large spores, 20x1 i/x, are probably wrongly 

nudipes Fr. — most likely to be H. Iongicaudum. 
punctatum Fr. — perhaps a form of Flammula lenta. 

radkatum (Cke.) Maire — a very uncertain species based on Cke. III. 459 (416). Maire thought 
the figures might be a slender H. senescens. Boudier suggested Flammula lenta. 
senescens (B2Lt&ch) F. sinuosum. 

sinapizans (Paul.) Fr. — ^left in but may be the same as H. sinuosum or a luxuriant form of H. 

truncatum Schaeff, — usually identified with Trickoloma (Rhodopaxillus) trnncatum but Lange 
maintains that there is a genSuine Hebeloma truncatum. 


LubricaeFx. ‘ Snbsiccae 

Flammula lenta (Pers.) Fr. Flammula ochrochlora Fr. 

lubrica (Pers.) Fr. astragalina Fr. 

gummosa (Lasch) Qudl. ^ 

spumosa Fr. var. salicicola Fr. 

carbonaria Fr, apicrea Fr, 

scamba (Fr.) Sacc. austera Fr. 

Indusiata Fr. 

Hebeloma fastibile (Pers.) Fr, 
versipelle Fr. 
mesophaeum (Pers.) Fr. 
testaceum (Batsch) Fr. 
strophosum Fr. 
subcollariatum B. & Br. 


Transactions British Mycological Society 

Subsiccae Lange {continued) 
Flammula fusus (Batsch) Fr. 
azyma Fr. 

flavida (Schaeff.) Fr. 
inaurata W. G. Sm. 
connissans Fr. 

Aldridgei Mass, 
clitopila Gke. 
decipiens W.G.Sm. 

{Fulmdula Romagnesi) 

Flammula hybrida Fr. 
limulata Fr. 
liquiritiae (Pers.) Fr. 
sapinea Fr. 
picrea Fr. 
flava (Bres.) Lange 


Flammula Aldridgei Mass.— left in, but this orange-coloured with the large spores 

requires confirmation, 
rflrwojfl Mass. — needs further elucidation. 

clitopila Gke. & Sm.— not known but W. G. Smith’s figure in Gke. Ill 468 (500) is so distinct 
that it is best left in list. Killerman reported it from Bavaria. 
decipiens W.G.Sm. — like the last, not well known, but reported by Killerman from Bavaria. 
decussata Vt. — ^no British record of this species. 

filia Fr. — not in Hym, Eur. Gke. III. 476 (432) may be a large form of Cortinarius rubicundulus. 
filicea Gke.— an alien on tree-fem stem. 
fioccifera B. & Br. = small form of Flammula lenta. 

gymnopodia (Bull.) Yx.-=Clitocybe tabescens sec. Quelet, but Gke. III. 465 (431) is a luxuriant form 
oI Flammula sapinea gyoymig on %2mdmt. 
helomorpha ¥T.=Ripartites tricholoma. 

inopus Fr. — no definite British record for this. Gke. III. 484 (446) is not this species, but 
Hypholoma radicosum Lange. 
jun€ina Sm. — F. lubrica ox ioxm oi F. alnicola. 
lupvna Fr.— doubtful record. 

mixta 'Bv. — Gke. ///. 471 {^^/^^^oithox F. carbonaria ox F. lubrka. 

nitens Gke. & Mass. — an ambiguous species. Massee’s figure in Gke. III. 1168 Ci 154) does not 
look like a Flammula. 

penetrans Fr.=F. sapinea Fr. Syst. Myc., where Fries cites penetrans as a synonym. 
purpurata Gke. & Mass. — exotic, on tree fern. 
ruhicundula Cortinarius {Ino.) rubicundulus. 
strigoc€ps 'Fx.=Ripartites tricholoma. 

trickoloma (A. & S.) Fr.— this agaric has been put in many genera and has (we hope) at last 
found a permanent home in the genus Ripartifes. 
vinosa (Bull.) Fr. — an ambiguous species. Gke. III. 466 (437) looks xaXhox liko Pleurotus Eryngii^ 
not yet recorded for Britain. 


(i) (Fayod) Heim 

Naucoria semi-orbicularis (Bull.) Fr. 
pediades Fr. 
vervacti Fr. 
arvalis Fr. 

var. tuberigena Opel, 
tabacina Fr. 

(2) (PhaeocollybiaHdm) 
Naucoria lugubris Fr. 
festiva Fr. 
cidaris Fr. 

(3) (Alnicola Kilhiier) 
Naucoria escharoides Fr. 

submelinoides Kiihner 
var. alnetorum Make 
Bohemica Vel. 
amarescens Quel. 

(4) {Macrocystis JELeirn) 
Naucoria cucumis (Pers.) Fr. 

(5) {Galerina Kiihner) 
Naucoria camerina Fr. 

sideroides (Bull.) Fr. 
triscopa Fr, 


Revised List of British Agarics and Boleti 

(6) Eu-naucoria Heim 
{a) Nudae 

Naucoria centunculus Fr. 
cerodes Fr. . 
horizontalis (Buii.) Fr. 
myosotis Fr. 
temuienta Fr. 

iff) Vestitae 

Naucoria carpophila Fr. 
efFugiens Qjuei. 
siparia Fr. 

(7) {Phaeomarasmius Scherff.) 
Naucoria erinacea Fr. 
rimulincola (Lasch) Rabh. 


Jfaucoria abstrma Fr.^^Flammula abstrusa sec. Lange, but not known if British record refers to the 
same species. 

anguinea Fr. — Gke. Ill 494 (455) appears to be copied from Fries. British record requires 

badipes Fr. — transferred to Galera. 

conspersa (Pers.) Fr. — ^has many interpretations. Lange places it in the Alnicola section. 
Romagnesi says it is the commonest Tubaria. 

echinospora W.G.Sm. — possibly a Leptonia from the rough spore, but it is an exotic as it grew 
in greenhouse. 

glandiformis Gke. — III 500 (490) B is probably a Cortimrius, 
graminicola (Nees) Fr. — probably Crinipellis stipitarius, 
hamadryas Fr. — ^may be JV. arvalis. 
hydrophilayL2iSS.=:Psilocyhe semi 4 anceata. 
innocua (Lasch) Fr. — Gke. Ill 498 (4^^) =: Flammula carbonaria, 
latissima Gke. Ill ^10 {4B2) — inadequately described. 

melinoides Fr. — ^what is usually called by this name in Britain is Galera mniophila, 
nasuta FisLlch, — Gke, III 11^^ {ii’]2)^Psilocybe semidanceata. 

nucea (Bolt.) Fr.— not known. Bolton’s figure in Gke. Ill 500 (490) suggests a form of PsatkyreUa 

Gke. & Mass. — Psilocybe sarcocephala. 

porriginosa Fr. — Gke. Ill 51 1 (510) may be a form of Tubaria furfuracea or Cortinarius scandens. 
pusiola Fr, — possibly Tubaria crobulus. 

feducta Fr. — best left out for further investigation; often confused with N, centunculus, 
fuhricataB, Sr.^v,'=^Marasmius ramealis. 

■scolecina Fr. — not really known; appears to belong to the Alnicola group. 

scorpiodes Fr. — ^recorded by Plowright, but there is some doubt what species it refers to. 

N. scorpiodes semnluOxi^e^N, Bohernica. 
semiflexa B. & Br. — may be Tubaria crobulus, 

semi-orbicularis (Bull.) Fr. — left in list but considered by many authors to be the same as 
JV. pediades which would have priority. 
sobria Fr. — ^probably a Tubaria, 
striaepes Gke, ^Galera pygmeo-affinis sec, Kflhjier, 

subglobosa (A. & S.) Fr.= JV. semi-orbicularis sec. Quelet, but Ricken describes something quite 

subtemulenta Lamb. — ^inadequate diagnosis. 

tenax fr, — ^identity doubtful. Gke. Ill 504 (617) is like a Tubaria and so is Ricken’s interpre- 
tation with its 4-angled heart-shaped spore 5-6 x 4-5/^. Rea describes a species growing in 
bogs on the stems of Potentilla Comarum and with spore is-ifi x 7-8 pt. This must be Naucoria 

Wieslandrii Fr.~— description inadequate. 

Gen. TUBARIA Gillet 

Tubaria furfuracea (Pers.) Gillet 
pellucida (Bull. ex. Fr.) Gillet 
trigonophylla (Lasch ex Fr.) Gke. 
anthracophila Karst, 
autochtona (B. & Br.) Sacc. 

Tubaria crobulus (Fr.) Karst, 
inquilina (Fr.) Gillet 

{Galerina Kiihner) 
Tubaria paludosa (Fr.) Karst, 
stagnina (Fr.) Gillet 

176 Transactions British Mycological Society 


tuharia cupdarU [hull.) sec. Quflet and Bresadola, but Cke. 7 Z/. 526 (60a) 

^T, pellucidaox T.furfuracea, 

mhda Fr.— best left out till British specimens are more clearly defined. 

mmcorum (Hoffm.) Fr. — unknown. ■ . , ^ . , . . . • u, , .r 

pellucida (Bull.) Fr.— this common species is left m, but it is questionable whether it differs 

from T. furfuracea. 

(Conocyhe Fayed) 

Galera spicula (Lasch) Fr. 
tenera (Schaeff.) Fr. 
antipus (Lasch) Fr. 
pilosella (Pers.) Fr. 
siliginea Fr. 
lactea Lange 
pygmaeo-affinis Fr. 

Mairei Kiihner 
plicatella Peck 

Gen. GALERA Fr. 

Galera appendiculata Lange & Kuhner 
campanulata Mass. 

{Galerina Earle) 

Galera graminea Vel. 
sphagnorum (Pers.) Fr. 
mycenopsis Fr. sensu Ricken 
mniophila (Lasch) Fr. 

Hypnorum (Batsch) Fr. 
rubiginosa (Pers.) Fr. 
nana (Petri) Kuhner 


Galera apalaVt . — a much disputed species. Set note on G, lateritia, 
campanulata Mass.— not known, but its strong smell and subfusoid spores are distinct characters, 
so it is not withdrawn. Massee cites Cooke’s figures 1174 (1156) for this species. 
confertai'^olt.) 'Fx.^G. antipm, 
flexipes KdiXSt. — unknown. 

iateritia Fr. — by many authors, including Cooke, Ricken and Kuhner, this epithet is used for 
a white or cream-coloured Galera with a bulbous base. The Friesian species refers to a species 
with a brick-coloured cap. Lange produced a new name for it, G. lactea, which we have 
adopted. It appears to be identical with Bolbitius tener Berk. But for its large size Galera 
apala Fr. would be the correct epithet, as the description fits closely. 
minuta Qneh — not known. 

mycenopsis Bx,— not known as described by Fries, and generally adopted for a fairly common 
Galera as described under this name by Ricken. 
ovalis Fr. — some authors depict this as a large form of G. tenera, but the G. ovalis of Fries has 
a fugacious ring and is not clearly defined. 

Fr.— cannot trace a British record. 
ravidaVx.-—Qke, III. ^2^ xni^the Pluteolus aleuriatus, 

Sahleri Quel.— seems little more than a form of G, hypnorum with conical pileus. 
sparteaYx . — seems to be a form of G. 

vestita Fr. — ^was reported by Bucknall in 1881, but it is doubtful if it corresponds to modern 
views of the species. 

vittaeformis Fr. = G. rubiginosa sec. Kiiliner. 


Crepidotus mollis (Schaeff.) Fr. 
calolepis Fr, 
alveolus (Lasch) Fr. 
applanatus (Pers.) Fr. 
fragilis Joss. 

Crepidotus variabilis (Pers.) Fr. 
sphaerosporus (Pat.) Lange 

{Phyllotopsis Gilbert & Donk) 
Crepidotus nidulans (Pers.) Quel. 


Bolbitius titubans (Bull.) Fr. 

Bolbitius vitellinus (Pers.) Fr* 
fragilis (Linn.) Fr. 


Revised List of British Agarics and Boleti 


Crepidotus chimonophilus B. & Br. — hardly different from C, variahilts, 
epibryus Fr.— if this differs from C. variabilis it requires clearer definition. 
epigaeus (Pers.) B. & Br. — ^Berkeley’s figure in Cke. III. 537 (516) indicates nodulose spores 
and may be a species of Claudopus. 

haustellaris 'F t.— C kQ. III. 536 (575) is from foreign specimens. The two old British records 
require confirmation. 

luteolus Lamb. — no authentic British record. 

Pmsoiii Pat. — ^British record appears to be a mistake. 
pezkoides (Nees) Fr. — diagnosis inadequate. 

Phillipsii B. & Br. — probably old C. variabilis. 
proboscideus Fr. — inadequately described. 

putrigenus B. & Curt. — an American species probably listed in error. 

Ralfsit B. & Br. — ^may be form of C. calolepis. 

versuius Peck — doubtful record. 

Bolbitius affinis Mass. — not known, but may be Galera lateriiia Fr.; more clearly defined in 
G. lactea Lange. 

apkalis W. G. Sm.-— doubtful if a Bolbitius or a Galera. 

Boltonii (FtXB.) Fr.^B. vitellinus. 

bulbillosus Fr. — a doubtful species reported from Scotland. 
flavidus (Bolt.) Mass.=J?. vitellinus. 

fragilis (Linn.) Fr. — ^left in but hardly distinct from B. vitellinus. 

grandiusculus Ckt. form B. vitellinus. 

niveus Mass. — from a palm house. May be Galera apala. 

rivulosus B. & Br. — in orchid house but probably only a form of B. vitellinus. 

tener Berk. = Galera lactea. ' 

titubans (Bull.) Fr. — left in but doubtful if different from B. vitellinus. 


{Myxacium Fr.) 

Cdrtinarius mucosus (Bull.) Fr. 
collinitus (Pers.) Fr. 
stillatitius Fr. 
elatior Fr. 
salor Fr, 
delibutus Fr. 
iliibatus Fr. 
vibratilis Fr. 
pluvius Fr. 
epipolius Fr- 
croceo-caeruleus Fr, 
Imdo-ochraeeus Berk. 

(Fhlegmacium Fr.) 

A. Scauri Fr. 

Gortinarius multiformis Fr. 
rapaceus Fr. 
intermedins Rea 
napus Fr. 
allutus (Seer.) Fr. 
purpurascens Fr. 

var. subpurpurascens Fr. 
caerulescens (Schaeff.) Fr. 

Gortinarius caesiocyaneus Britz. 
dibaphus Fr. 

var. xanthophylius Cke. 
glaucopus (Schaeff.) Fr. 
herpeticus Fr. 
cyanopus (Seer.) Fr. 
calochrous (Pers.) Fr. 
fulgens (A, & S.) Fr. 
fulmineus Fr. 
elegantior Fr. 
flavescens (Cke.) Henry 
turbinatus (Bull.) Fr. 
lutescens (Rea) Henry 
prasinus (Schaeff.) Fr. 
atro-virens (Kalch.) Fr. 
orichalceus (Batsch') Fr. 
rufo-olivaceus (Pers.) Fr. 
scaurus Fr. 

B, Cliduchi Fr. 

Cortinarius sebaceus Fr. 
claiicolor Fr. 
triumphans Fr. 
praestans (Cordier) Sacc. 
balteatus Fr. 

178 Transacticm British Mycological Society 

Clidmki ft, {continued) 

, Cortiiiarim largus Fr. ■ . 

var, nemorensis Fr. 
variicoior (Pers,) Fr. 
varius (Schaeflf.) Fr. 
cumatilis Fr. 
mfractus (Pers.) Fr. 
percomis Fr. 

Riederi (Weinm.) Fr. 
saginus Fr. 

G. Elastici Fr. 
Gortinarius turmalis Fr, 
cristailinus Fr. 
causticus Fr. 
emollitus Fr. 
decolorans Fr. 
decoloratus Fr. 
porphyropus (A. & S.) Fr. 
(croceo-caeruleus Fr.) ; see Myxacium 
papulosus Fr. 

{Inoloma Fr.) 

Gortinarius humicola (Quel.) Maire 
opimus Fr. 
turgidus Fr. 
argutus Fr. 
pholideus Fr. 
tophaceus Fr. non Qudl. 

Bulliardii (Pers.) Fr. 
bolaris (Pers.) Fr. 
rubicundulus (Rea) Pearson 
violaceus Fr. 
albo-violaceus (Pers.) Fr. 
suillus Fr. 

argentatus (Pers.) Fr. 
fusco-tinctus Rea 
cyanites Fr. 
muricinus Fr. 
bircmus Fr. 
traganus Fr. 

var. finitimus Weinm. 
vinosus Glee. 

{C, malachiusft,,^t^ Telamonia) 
(Dermocybe Fr.) 

A. Ochroleuci Konr. & Maubl. 

Gortinarius ochroleucus (Schaeff.) Fr. 
decumbens (Pers.) Fr. 
camunis Fr. 
tabularis (Bull.) Fr. 

B. Cinnamomei Konr. & Maubl. 

Gortinarius cinnamomeus Fr. 
croceo-conus Fr. 

Gortinarius semi-sanguineus Gillet 
sanguineus ( Wulf.) Fr. 
malicorius Fr. 
cinnabarinus Fr. 
anthracinus Fr, non Qu6L 
uliginosus Berk. 

Queletii Bat. 
phoeniceus (Bull.) Maire 
orellanus Fr. non Quel. 

G. Anomali Konr. & Maubl. 
Gortinarius anomalus Fr. 
iepidopus Gke. 
azureus Fr. 
caninus Fr. 
myrtillinus Fr. 
spilomeus Fr. 

D. Veneti Konr. & Maubl. 

Gortinarius cotoneus Fr. 
venetus Fr. 

raphanoides (Pers.) Fr, 


A. Platyphylli ft, 

Gortinarius macropus Fr. 
laniger Fr. 
bivelus Fr. 
licinipes Fr. 
bulbosus (Sow.) Fr. 
malachius (Fr.) Pearson 
bovinus (Seer.) Fr. 
triformis Fr. 
glandicolor Fr. 
brunneus (Pers.) Fr. 
biformis Fr. 
brunneo-fulvus Fr. 
armillatus Fr. 
limonius Fr. 
callisteus Fr. 
helvolus Fr. 
hinnuleus Fr. 
gentilis Fr. 
nitrosus Gke. 
helvelloides Fr. 
penicillatus Fr. 
torvus Fr, non Quel, 
scutulatus Fr. 
impennis Fr. 
evernius Fr. 
quadricolor Fr. 
plumiger Fr. 


Revised List of British Agarics and Boleti 

B. LeptophylliVi. 

Gortinarius paleaceus Fr. 
fiexipes Fr. 

hemitriciius (Pers.) Fr. 
rigidus (Scop.) Fr. 
stemmatus (Fr.) Ricken 
iliopodius (BuU.) Fr. 
incisus (Pers.) Fr. 

Cookei Quei. 

psammocephaius (Bull.) Fr. 
flabellus Fr. 

Iris Mass. 

ammophilus Pearson 

(Hydrocybe Fr.) 

A. Firmiores Fr. 

Gortinarius duracinus Fr. 
pseudo-duracinus Henry 
subferrugineus (Batsch) Fr. 
imbutus Fr. 
firmus Fr. 
armeniacus Fr. 
damascenus Fr. 
dilutus Fr. 
tortuosus Fr. 


Cortinarius albo^cyaneiis Fr. — ^not known. 
angulosus Ft.--v2Lnom\Y interpreted. Cke. ///. 1192 (1178) may be a form of C. cinmmomus 
or C. venetus. 

arenatml^x. TioiiQyS..^C. pholideus, 
arpinaceus 'Fr,=zC, mucosus. 

camphoratus Fr. — doubtful record; not the same as that of Ricken or Henry; Cke. IlL 751 (771) 
is probably C. or C. 

coruscansVx. — ^not known. Cke. III. 730 (733) may be C. sebaceus. 
colymbadinns Fr. — record doubtful. 

carrosus Fr. — Cke. IlL 715 (715) looks like a dwarf form of C. multiformis, 
croceofulvus fDC.) ^x.‘ — Cke. 7 //. 1191 (1193) suggests C. 

crassus Fr. — ^has several interpretations. Cke. ///. 685 (695) is more like C. praestans. 

crocolitus QyiS.. — hardly seems distinct from C. 

depressus Fr. — doubtful. Cke. III. 854 (860) looks like C. duracinus. 

detonsus Vx. — ^record by Stevenson not this species. 

diabolicus Fx. — Cke. ///. 765 (816) looks like C. 

dolobratus Fr. — ^not known what this is. Cke. ///. 845 (81 1) has been referred to several species. 
duracinus Fx. — ^Henry has established a new species C. pscudo-duradnus now included in British 
list since it is based on Cooke and Massee’s spore measurements 5 X3/X. Henry’s measure- 
ments are 5-5JX4I— 5 ft, so it is extremely doubtful if the two species are identical. 
fallaxQyxB.. — ^no British record can be traced. 
fulvescens Fr. — ^no British record can be traced. 

grallipes Fr. — Cke. III. 738 (734) by W. G. Smith looks like a large Flammula gummosa. The true 
grallipes has a prominent acute umbo. 
ianthipes — ^not known. 

illuminus Fr. — ^hardly known. Cke. III. 830 (841) suggests C. rubicundulus. 
impennis Fr.' var. lucorum Fr. — ^record is based on Gke. III. 1190 (1192) which looks more like 
C. evemius. 

Gortinarius saturninus Fr. 
bicolor Fr. 
castaneus (Bull.) Fr. 
balaustinus Fr. 
colus Fr. 

isabellinus (Batsch) Fr. 
renidens Fr. 
uraceus Fr. 
pateriformis Fr. 
rubricosus Fr. sensu Lange 
holophaeus Lange 

B. Tenuiores Fr, 

Gortinarius rigens Fr. 
leucopus (Bull.) Fr. 
Krombholzii Fr. 
obtusus Fr. 
scandens Fr. 
acutus (Pers.) Fr. 
erythrinus Fr. 
decipiens (Pers.) Fr. 
germanus Fr. 
saniosus Fr. 
fasciatus Fr. 

Junghuhnii Fr. 


i8o Transactions British Mycological Society 

Coniinarius infucatm Fr.— awaits more detailed description. 
injucundus (Weinm.) Fr. — Cke. Ill, Bog (823) is old C. largus. 

Iris Mass. — ^rather dubious, but not withdrawn. 

imgularis Fr.-— based on old record by Bolton which from original description and figure was 
Volvaria speciosa f. gloiocephala with remnant of volva. 
jubarinus Fr. — doubtful if distinct from C. cinnamomeus, 
latus (Fers.) Fr. — cannot trace British record. 

Ucinipes Fr. — ^left in because of Gke. Ill 792 (819) which may however be a form of C. bivelus, 
livido-ochraceus Berk. — left in with some doubt because the flesh is mild; otherwise W. G. 

Smith’s figures in Cke. Ill 739 (767) look like pale forms of C. croceo^caerulem. 
lustratus Fr. — Gke. Ill 688 (799) may be C. argentatus. 
mkrocydm Fr. — unknown species. Cke. Ill 793 (865) = C, glandicolor, 
milvinus Fr, — seems identical with C. raphanoides. 

muciflmts Fr. — Cke. Ill 735 (740) is typical C, collinitus. Most authors describe a form of 
C, datior under this name. 

nitidm (Schaeflf.) Fr. — based on Cke. Ill 1189 (1191) which may be C, delibutus. 

olivascens (Batsch) Fr. — ^remains to be seen if the British record is other than C. infractus, 

paragaudis Fr. — doubtful. 

periscelisVt,'=‘C, paleaceus. 

phrygianus Fr. — ^fuller description needed. 

plumiger Fr. — doubtful if British record refers to this very rare species. 

privignus Fr. — requires further investigation; at present it seems much the same as C. ochro- 

punctatus {^ets,) 'F t , — doubtful record. 

qmdricolor (Scop.) Fr. — left in because of the excellent figure in Cke. Ill 799 (867) but rather 

Reedii Berk. — ^not known what this refers to. Cke. Ill 848 (843) looks like an Inocybe. 
riculatusFx. — doubtful. 

Riederi Fr. — uncertain, but left in because of Epping record and Cooke’s excellent figures, 
694 (702). 

rubellns Gke. = C. orellanus Fr. 

russus Fr.-— doubtful. Cke. Ill 696 (751) looks more like C. sub-ferrugineus, 

Fr.-— doubtful if distinct from C. 
serariusFf, — record doubtful. 

sublanatus Sow.-C. pholideus. Cke. ///. 761 (762) is also pholideus. The C. suhlanatus of other 
mxthot$ k C, cotorieiis, 
submtatm Fr. =^ raphanoides 01 venetus, 
iestaeeus Cke,^C. rufo-olivaceus, 
turbinatusvzT. lutescensFe^—C. lutescens {FLeo) Fiemy, 
unimodus Fritz. — Gke. ///. 844 (859) = C. 

Fr,— record uncertain. 

valgus Fr.— unknown species. Gke, III 785 (750) may be C. raphanoides or C. venetus. 
vespertinus Fi, — ^not a convincing record. 

vinosus Cke.— has been queried. Cke. Ill 758 (759) looks rather like C. subpurpurascens but the 
spores are different and it may well be a distinct species. 
violaceo-fuscus Cke. k— Inocybe obscura. 


1 . Levisporae 
{a) Depauper ata 
Inocybe rhodiola Bres. 
Patouillardii Bres, 
Bongardii (Weinm.) Quel, 
cervicolor (Pers.) Karst, 
calamistrata Fr. 
hirsuta Lasch 

Inocybe relicina Fr. 
fastigiata (Schaeflf.) Fr. 
Cookei Bres. 
maculata Bond, 
perlata (Weinm.) Fr. 
perbrevis Fr. sensu Cke. 
dulcamara (A. & S.) Fr. 
squamata Lange 

Revised List of British Agarics and Boleti i8i 

{b) Muriculatae 

Inocybe pyriodora (Pars.) Fr. 
var. incarnata (Bres.) Maire 
corydalina Quel, 
iucifuga Karst, 
liaemacta Berk. & Gke. 

var. rubra Rea 
Godeyi Gillet 
bystrix Fr. 

mutica Fr. sensu Gke. 
cincinnata Fr. 
obscura (Pers.) Fr. 
griseo-liiacina Lange 
geophylla (Sow.) Fr. 

var. lilacina Fr. 
vatricosa Fr. 
flocculosa Berk, 
abjecta Karst, 
eutheles B. & Br. 
pallidipes Ell. & Ev. 
descissa Fr. 

var. brunneo-atra Heim 
auricoma (Batsch) Fr. 
deglubens Fr. 
postemla (Britz.) Lange 
Queletii Maire & Konr. 
sambucina Fr. 
brunnea Quel, 
lacera Fr. 
halophila Heim 
serotina Peck 

11 . Nodulisporae 

(Astrosporma Scbroet.) 

Inocybe asterospora Quel, 
napipes Lange 
umbrina Bres. 
striata Bres. 
margaritispora Berk, 
umboninata Peck 
trechispora Berk, 
grammata Quel, 
fulva Rea 
fibrosa Sow. 
duriuscula Rea 
praetervisa Quel, 
scabella Fr. sensu Cke. 
fulvella Bres. 

Boltonii Heim 
lanuginella Schroet. 
decipientoides Peck 
lanuginosa (Bull.) Fr. 
longicystis Atk. 
umbratica Quel, 
maritima Fr. 
fasciata Cke. & Mass. 

Rennyi B. & Br. 
petiginosa (Fr.) Gillet 

III. Verrucosisporae 
Inocybe calospora Quel. 


Inocybe abjecta Fr. — left in, but doubtful if other than a small form of /. flocculosa, 

Bresadolae grammata QnB., 

Bticknallii Nolanea fumosella. 

caesariata Fr. — Gke. III. 437 (388) probably /. hirsuta. 

carpta (Scop.) Fr,- — ^has difierent interpretations. Gke. III. 419 (426) is probably /. hirsuta, 
while Bres. Icon. 756 has nodulose spores. 

ClarkuB. 81 Br. = I. geophylla. 

conformata Karst.— seems to conform to /. obscura. 

destricta Fr. — not clearly defined; may he I.jurana Pat. which, though near to /. rkodiokj does 
not turn the rich claret colour so characteristic of the latter. But the synonymy of these two 
species is uncertain. 

duriuscula Rea — left in, though Kiihner thinks it a form of I. grammdta. 

fasciataXI!R&. & Mass.— retained but not known since first found in Kew Gardens; may be an 

Gailliardi Giilet=L calospora. 
infidaVtch^^I. umbratica QvSl. 
mamillaris Buss . — ^description insufficient. 

margaritispora Berk. — ^not known to modern mycologists but from Gke. ///. 432 (505) it appears 
to be a distinct species, unless considered as a large form of I. calospora. 
mimica M.USS. — probably /. 

mutica Fr.— not really known but Cke. ///. 418 {382) appears to represent a distinctive thing 
and the name is retained. 

i 82 Transactions British Mycological Society 

InGcybe nigro-disca Vtch--t€Cotd donh^^, 

phueocephala Gke.—the bright ferruginous spores indicate a Cortinarius. 

pgfhnvis (Weinm.) Fr, — left in since Heim accepts it on basis of Cke. Ill, 434 (519) though this 
plate does not look a bit like an 

plumosa Bolt. — doubtful if an Inocybe, The /. plumosa of Konr. & Maubl. is I, lanuginosa. 
prQximeUaK.BXSt,=I. asterospora, 
psmdo^fastigiata^tdL^I. fastigiata. 

Mennyi B. & Br. — ^left in with some doubt. Heim examined type specimen, and accepts species, 
rimosa (Bull.) Fr. — this epithet has been applied to many different species and has been aban- 
doned. The rimosa of Fries was probably I, asterospora; the description in Syst. Myc. certainly 
suggests this. 

Sabuietorum B. Sl Curt =L lanuginosa, 

seabra (Mull.) Fr. — epithet used in different senses; uncertain what British record refers to. 

Cke. III. 413 (391) is cited by Heim as near I, splendens. 
schista Cke. & Sm.^ — description inadequate. Cke. Ill, 423 (504) looks more like a Cortinarius, 
sindonia Fr.^ — ^hardly distinct from I, geophylla, 
squarrosa I, cincinnata, 

Trinii Weinm.— uncertain what this refers to; may be a small form of I, Patouillardii. 
tomentosa Qxii€l. (1888)=/. eutheles B. & Br. (1865). The I, tomentosa of Junghuhn (1830) is 
doubtful and has nothing to connect it with /. eutheles. 
mirkosa Fr. — ^not well known; usually on wood and retained for this reason but clearer de- 
finition needed. 

violaceo-folia Peck = a Cortinarius sec. Heim. 

Whiiei B. & Br.=/. vatricosa Fr. sec. Heim. 


Paxillus involutus (Batsch) Fr. ( Tapinella Gilbert) 

atrotomentosus (Batsch) Fr. Paxillus panuoides Fr. 

(See also among the Boletales) 


Paxillus Alexandra 'Fr. = Cliiocybe Alexandra (Gillet) Konrad, but there is no clear British record. 
The following extract from a letter written by G. Massee to A. A. Pearson, dated 19 Jan. 
1916, disposes of one record: ‘Cooke and I were responsible for the record, but in reality 
it was founded on a small specimen of the old Lactarius exsuccusi 
cfossus Fr. — Cke. III. 864 (861) is probably old Phylloporus rhodoxanthMS. 
extenuatus Bx. — Cke. ///. 863 (873) looks like amam. 

giganteus {Sow.) Px.^Clitocybe gigantea, 

lepista Fr.— has different meanings; sensu Ricken it is Clitocybe {Rhodoxanthus) mundula; sensu 
Cooke it may be Clitocybe amara. 

leptopus Px, — said to be confined to alders, but hardly distinct from P. involutus. 

lividus Cke.^ — ^probably Tricholoma cinerascens. 

orcelloides Cko. M.Q.SS, — mdi.y ho Clitocybe fallcix, 

panaeolus Px.^ Ripartites tricholoma. 

paradoxus (Pidlch.) Q\io\.s=zPfylloporus rhodoxanthxis. 

porosm BtTls..'~--yrQhQhly Phylloporus rhodoxanthus. 

revolutus Cke. — ^Gooke’^s figure 865 (862) looks like Hygrophorus cinereus but round spores, 
if correct, would not fit, 


I. MundaePt. Stropharia coronilla (Bull.) Fr. 

Stropharia aeruginosa (Curtis) Fr. hypsipus Fr. sensu Lange 

albo-cyanea (Desm.) Fr. melanosperma (Bull.) Quel. 

Worthingtonii Fr. squamosa (Pers.) Fr. 

inuncta Fr. var. thrausta Kalch. 

luteomtem (Vahl) 

Revised List of British Agarics ar^ Boleti 183 

II. Merdariae Fr. 

Stropharia semi-giobata (Batsch) Fr. 
merdaria Fr. 

III. Hypholomoideae 
Stropharia caput-Medusae Fr. 
scobinacea Fr. 

Jerdonii B. & Br, 

Stropharia spintrigera Fr. 

Hornemannii Fr. 

Percevalii B. & Br. 

Ferrii Bres. 

IV. Psathyroideae Lange 
Stropharia psathyroides Lange 

(See also Anellatia and Psathyra pennata) 


Stropharia Batarrae Fr. — ^no British collection can be traced; it is a problematical species. 
coturnata Fr. — description inadequate. 
depilata Hornemannii^x, Syst. Myc. 

hypsipus Fr. — Massee’s figure in Cke. III. 571 (619) is problematical, but .S', hypsipus Fr. sensu 
Lange has been collected and is retained in list. 
luteo-nilens (Vahl) Fr. — ^left in, but it is doubtful if Cke. 111 . 564 (604) represents this species. 
inelasperma Pt.^melanosperrna stc.BullmxdiS 

obturata Fx.^S. coronilla. 

Percevalii B. & Br. — ^left in list though much like Hornemannii except that spores are much larger 
sec. Cke. Ill 554 (550). 

punctulata (Kalch.) Fr. — a problematical species. 

rugoso-amulata Farlow (1929). This has now been identified with S. Ferrii Bres. (1928), a species 
first found in northern Italy and recently reported from Switzerland. The descriptions do 
not exactly tally, but in view of the remarkable variability observed in the British specimens, 
we must conclude that they represent the same species. 
squamulosa Mass. = < 5 '. aeruginosa, 
stercoraria Fr.=*S'. semi-globata. 
versicolor (With.) Fr.^ — insufficiently described. 


{Nematoloma Karst.) Hypholoma Candolleanum Fr. 

Hypholoma fasciculare (Huds.) Fr. cascum Fr. 

sublateritium Fr. chondrodermum (B. & Br.) Lange 

var. squamosum Cke. leucotephrum B. & Br. 

capnoides Fr. lacrymabundum Fr. non Bull, 

radicosum Lange 

dispersum Fr. {Laeririiaria 'PaX,) 

{Drosophila Qp^.) Hypholoma velutinum (Pers.) Fr. 

Hypholoma hydrophilum (Bull.) Fr. pyrotrichum (Holmsk.) Fr. 

catarium Fr. sensu Cke. 


Hypholoma aellopum Ft. — description inadequate. 

appendiculatum Fr, — an ambiguous species. The agaric usually given this name is H, Candolleanum, 
catarium Fr. — retained but may only be small H, hydrophilum, 
egenulumB. Sc Bx.^^H, Candolleanum. 
elaeodes¥T.=H. fasciculare. 

epixanthumFx,' — ^probably a form ofH. capnoides sec, Seth Lundell. The British records usually 
refer to young specimens of 
incomptum Mass, — ptohahly Stropharia Horriemannii, 
instratum H . chondrodermum. 

irroratumFiaxst. — doubtful, but suggests H. 
lanaripes Candolleanum. 

melantinum Fx.-==^ Stropharia scobinacea Ft, stm\xBSck^xk. 

oedipus Cke. — not known, but Cke. 579 (587) showing a glutinous dark-coloured agaric is 
characteristic enough to keep in list. 
silaceum (Pers.) Fr. — ^not sufiSciently described. 

184 Transactions British Mycological Society 


{Drosophila Quth) 
Psiiocybe sarcocephala Fr. 
spadicea Fr. 
ammopliila Mont, 
clivensis B. & Br. 
cernua ( VaM) Fr. 
canofaciens Cke. 
catervata Mass. 

{Hypholoma Fr. em. Kuhner) 
{Mematoloma Karst.) 
Psiiocybe ericaea (Pers.) Fr. 
sub-ericaea Fr. 
uda (Pers.) Fr. 

Polytricbi Fr. 
elongata Fr. 

{Deconica W. G. Sm.) 

Psiiocybe coprophila (Bull.) Fr. 
bullacea (Bull.) Fr. 
physaloides (Bull.) Lasch 
atrorufa (Schaeff.) Fr. 
cyanescens Wakef. 

{Panaeolina Maire) 
Psiiocybe foenisecii (Pers.) Fr. 


Psiiocybe semi-lanceata Fr. 
var. caeruiescens Cke. 


Psiiocybe agraria Fr. — description incomplete. Cke. III. 597 (622) looks like a Mycena. " 
areolata (Klotsch) Berk.— Cke, III. 596 (570) may be a sundried Hypholoma capnoides. Cke. Ill 
1182 (1177) also was given this name but Massee referred it to P. virescens. 
airo-brunnea (Lasch) Fr.— identity uncertain and no British record known. 
callosa Fr. — ^has many interpretations. 
canobrunnea (Batsch) Fr.— identity of British record uncertain. 
canofaciens Cke. — ^left in, but not known to modern authors. 
catervata Mass. — ^not known but left in, as it is fully described. 
cfu>ndrodennal&. Hypholoma clwndroderma. 

compta Fr. — Cke. III. 603 (589) is probably a Galera. 

cyanescens (1946)— published in these Transactions, xxix, 141. In a footnote on 

p. 1 65 it was pointed out, that it might prove to be the same as Hypholoma cyanescens Maire 
(1928) as further described and figured by Malen^on in 1942. Comparison with the un- 
published drawings of Miss Wakefield shows marked differences. The shape of the pileus 
is very different; so is the attachment of the gills. For the present they are best considered 
as distinct species. 

hebes Fr. — best withdrawn till further elucidated. 

helvola (Schaeff.) Mass. — seems to be a Panaeolus. 

nemophila Fr,— inadequately described and no British record found. 

nuciseda'Ex. — Cke. ///. 601 (609) is a 

sarcocephala Fr. var. Cookei Sacc.=large form of P. spadicea. 

scobicola B. & Br.— Cke. III. 598 (607) looks like Hypholoma Candolleanum. 

squalens Fr. — ^not known; description insufficient. 

iegularis (Schum.) Fr. — not known; and no British record found. 

virescm (Cke. & Mass.) Mass.-referred to Cke. III. 1182 (1177) over the name P. areolata; 
all very doubtful. 

Gen. PSATHYRJELLA Fr. {Drosophila Quel. 

I. (PsathyraPT.) PannucialLoxst. 

Veil evident 

Psathyrella Gordonii (B. & Br.) comb. nov. 
gossypina (Bull, ex Fr.) comb. nov. 
fibrillosa (Pers. ex Fr.) comb, nov; 
bifrons (Berk.) A. H. Smith 
semi-vestita (Berk.) A. H. Smith 
pennata (Fr.) comb. nov. 

Psathyrella pennata f, annulata Pearson 
nolitangere (Fr.) comb. nov. 
frustulenta (Fr.) A. H. Smith 

II. Eu-Psathyralsaxigc 
Margin naked. Spores small 
Psathyrellaspadicea-grisea (Schaeff. exFr.) 
A. H. Smith 

obtusata (Fr. sensu Lange) A. H. Smith 


Revised List of British Agarics and Boleti * 185 

IIL Psathyrella Fr. 

Margin naked. Spores large 
Psathyrella caudata Fr. 
conopilea Fr. 
subatrata (Batsch) Fr. 
gracilis Fr. 

var. corrugis (Pers.) Lange 


Psathyra Fr.- — This generic epithet is withdrawn because it is preoccupied by a genus of flowering 
plants, as pointed out by Kuhner in Bull. Soc. mycoL Fr. m, 1 1 . There being no clear line of 
demarcation, it is desirable to combine in one genus the species that hitherto have been 
divided between Psathyra and Psathyrella; the latter epithet can be adopted, though some 
authors wish to adopt Drosophila Quel. 

Psathyra elata yLsLSS. = Psatlyrella conopilea. 

fatua Fr. — ^it is not clear what this refers to and is best left out for clearer diagnosis. 
glareosa B. & Br. — Gke. III. 610 (591) looks like Mycena zephira. 
gyrqflexa Fr. — ^wants clearer definition. Gke. III. 1184 (970) is more like P. hiascens. 
helobia Kalch. — doubtful record. 

Lascosii Rabenh. — description insufficient. 

mastigera B. & Br. — hardly distinct from Psathyrella caudata. 

microrhiza (Lasch) Fr. — Gke. III. 622 (596) is Psathyrella caudata. 

neglecta Mass. — ^still neglected and unknown. 

pellosperma (Bull.) B. & Br. — definition incomplete. 

tenuicula Karst. — ^identity doubtful. 

Psathyrella arata Berk.— Gke. III. 63 7 (636) is hardly distinct from conopilea though unusually sulcate. 
empyreumatica B. & Br, — description inadequate. 

hiascens Fr. — ^referred to Coprinus by some authors but the gills do not deliquesce. It is question- 
able whether it is distinct from P. Fr. 

hydrophora (Bull.) Fr. — doubtful. 

trepida Fr. — not certain to what the British record refers. 


{b) Nudi 

Panaeolus acuminatus (Schaeff.) Fr. 
fimicola Fr, 
subbaiteatus B. & Br. 

Gen. ANELLARIA Karst. 

Annellaria semi-ovata (Sow. ex Fr.) Annellaria fiimiputris (Fr.) Karst, 

comb. nov. 


Panaeolus caliginosus Qun^.) Ft. =P. acuminatus. 

egregitis^Qk&. III. {62^ macy he Hypholoma velutinum or H. pyrotrichum, 
leucophanes B. & Br. Gke. ///. 625 (927) is probably a form of Anellaria semi-ovata. Ricken 
describes a different species under this name. 
phalaenarumPv.— Anellaria semi-ovata vdihout ting. 

Anellaria separata (Juixsa.) A. semi-ovata. In the Syst. Myc. thh appears in the section 

Coprinarius as Agaricus semi-ovatm, although separatus Linn, is given as a synonym. Sub- 
sequently Fries adopted the Linnaen name, but by the rules we cannot avoid using semi- 
ovata. The Agaricus semi-ovatus on PI. i$i of Sowerby is clearly this species. 
scitula Mass.— a small species of Coprinus with a peronate ring. As it was growing in a pot it 
was probably an exotic. 


{a) Appendiculati 

Panaeolus papilionaceus (Bull.) Fr. 
retirugis Fr. 

campanulatus (Linn.) Fr. 
var. sphinctrinus (Fr.) Bres. 

Psathyrella prona Fr. 
atomata Fr. 
typhae Kalch. 
crenata Lasch 

IV. [Pseudocoprinus Kuhner) 
Psathyrella disseminata (Pers.) Fr. 
hiascens Fr. 

1 86 Transactions British Mycological Society 



Psalliota augusta Fr. 

Elvensls B. & Br. 
arvensis (Schaeff.) Fr. 
silvicola (Vitt.) Sacc. 
viilatica Brond. sensu Bres. 
xanthoderma Gene V. 
var. lepiotoides Maire 
var. obscurata Maire 
var. grisea Pearson 
comtula Fr. 
amethystina Qtk^L 
mssiophylla (Lasch) Fr. 
dulciduia Schulz, 
pallens (Lange) Rea 
rubella (Giilet) Rea 

(2) Rufescentes J, Schaeffer 
{a) Gill edge fertile 
Psalliota campestris (Linn.) Fr. 

{b) Gill edge sterile 
Psalliota hortensis Gke. 
vaporaria (Vitt.) J. Schaeffer 
silvatica (Schaeff.) Fr. 
iitoralis Wakef. & Pearson 
arenicola Wakef. & Pearson 

(3) iSanguinolentes J. Schaeffer 
Psalliota sanguinaria Lange 
haemorrhoidaria Karst, 
exserta Rea non Viv. 

(4) Immutabiles 
Psalliota flocculosa Rea 
impudica Rea v 


Psalliota, — We have left out the numerous varieties of P. campestris and P. arvensis because it is 
not known with any certainty to which species they should be attached. On the other hand 
we have left in all the satellites of P. comtula though we have a suspicion that they could 
all be reduced to one species: P- amethystina. 

Psalliota exserta Viv. — under this epithet Rea has described a bleeding mushroom which is not 
the exserta of Viviani, a non-bleeding species, though the original figures are misleading. 
Viviani states: sua mantien bianca esposta alV aria.' We have gathered Rea*s 

species on chalk downs in Sussex but it requires further study before re-naming. 
fiavescens GiWti^i^P. silvicola. 
jSierrara Schulz. =P. augusta. 
peronaia'M.2t.^.=P. augusta. 

praiensis (Schaeff.) Fr. — ^identity uncertain — ^not unlikely to be one of the varieties of 
P. xanthoderma. 

sagata Fr.— Gke. III. 1177 ( 9 ^ 8 ) may be a large P. rubella. 

setigera Fr.-— seems to have slipped in the British list by mistake. 

subgibbosa Fr, — description insufficient. Cke. 111 . 551 (552) is Pholiota aegerita. 

Pilosace Algeriensis Pr, — doubtful species; probably a Psalliota with ring rubbed off. 

Ckrkeinda O. Kuntz {Chitonia Fr.) = C. rubrkeps (Gke. & Mass.) Rea. An alien species on soil in 
Aroid house. 


I. Pelliculosi Fr. 

(a) Jnnwto* Lange 

Goprinus comatus {FL Dan.) Fr. 
ovatus (Schaeff.) Fr. 
sterquilinus Fr. 

ijb) AiramentariiPu 

Goprinus atramentarius (Bull.) Fr. 
fuscescens (Schaeff.) Fr. 

ip) ToimntosiPx. 

Goprinus picaceus (Bull.) Fr. 
similis B. & Br. 
lagopus Fr. 
radiatus (Bolt.) Fr. 
lagopides Karst, 
macrocephalus Berk, 
cinereus (Schaeff.) Fr. 
naacrorhizus (Pers.) Rea 
echinosporus Buffer 

Revised List of British Agarics and Boleti 

(d) Phaeospori Lange Goprinus stercorarius (Bull.) Fr. 

Coprinus domesticus (Pers.) Fr. tigrinellus Boud. 

urticaecola (B. & Br.) Buller stellaris Qu^. 

FriesiiQuel. velox Godey 

filiformis B. & Br. 

II. Farinosi Lange curtus Kalch. 

radians (Desm.) Fr. 

{a) Annulati micaceus (Bull.) Fr. 

Goprinus ephemeroides (Bull.) Fr. 

Hendersonii Berk. HI. Lange 


(b) Exanmlati 

Goprinus cordisporus Gibbs 
Patouillardii Qu^L 
niveus (Pers.) Fr. 
dilectus Fr. non Lange 
roseotinctus Rea 
narcoticus (Batsch) Fr. 

Goprinus tergiversans Fr. sensu Ricken 
tardus Karst, 
ephemerus (Bull.) Fr. 

f. bisporus (Lange) Joss, 
congregatus (Bull.) Fr, 

Boudieri Quel, 
hemerobius Fr. 
plicatiiis (Curt.) Fr. 


Coprinus alternatus (Schum.) Fr. — not clearly defined. 
apthosus Fr. — no British record. Gke. III. 653 (666) is a Hypholoma. 
aqmtilis Peck — ^insufficiently described. 
araius'B. %L'Bv.-=\ 3 Xg€i C. micaceus. 

Bresadolae Schulz. — doubtful record. 
bulbillostis V Sit. ^C. ephemeroides. 
cothurnatus Godey — awaits fuller description. 

cordisporus Gibbs — left in but may prove to be the same as C. Patouillardii. 
deliquescens (Bull.) Fr. — not known to what this refers. 
digitalis Px. — ^needs confirmation. 

^^Mrw^zAy Qud.— -doubtful if a Coprinus. The violet almond-shaped spore suggests a white form 
of Psilocybe coprophila. 

erythrocephalus (Lev.) Fr. = C. dilectus sensu Lange non Fr. British record is doubtful. 
exiinctorius (Bull.) Fr. ~ C. micaceus. 

filiformis B. & Br. — ^left in on the strength of the spore measurements by Massee 5 x 4/x, but 
a doubtful species. 

fimetarius (Linn.) Fr.=C'. cinereus. The latter name was used in Syst. Myc. and the former cited 
as synonym. 

flocculostts TiG . — problematical species; perhaps not a 

frustulosum Sacc. — ^hardly different from C. roseo-tinctus which is retained, though it may be 
identified with C. 

Gt^teTMass. & Grossl.=a form of C. 

hemerobius Fr. — ^left in for further observation, but there is some doubt about its identity. It 
seems to be a large C. plicatiiis with an elliptical spore, but this may only be the side view 
of die plicatiiis 

Fr.— -doubtful to what this refers. 
oblecttts (fio\t.) Px. = C. sterquilinm sec.Bvller. 

papillatus (Batsch) Fr. — requires further elucidation; may only be C. plicatiiis. 

platypus Berk. — an alien. 


Schroeteri Karst. — ^not known; no British record found. 

Speggazinii K2txst. — an exotic. 

SpragueiB. & Curt. — not 2i Coprinus. 
squamosus M-oxg. — doubtful if distinct from 

i88 Transactions British Mycological Society 

Coprinm trumofum {Sch.Q.&'S,) miccuieus, 

tuberosus Quel. = C. stercorarius. 

tomentosm (Bull.) Fr. — another problematical species. To Ricken it seems little different from 
C. m warm but the Bulliard figure is not unlike C. 
umbrinus Cke. & Mass. — ^hardly differs from C. sterquilinus. 
mhaceo-mimmus Grossl. — seems identical with C. HendersoniL 

Gen. GOMPfflDIUS Fr. 

Gomphidius rutilus (Schaeff.) Fr. Gomphidius gracilis B. & Br. 

(=:viscidus) glutinosus (Schaeff.) Fr. 

maculatus (Scop.) Fr. roseus Fr. 



Strobilomyces strobrlaceus (Scop.) Berk. 

Porphyrellus porphyrosporus (Fr.) Gilb. 


Gyroporus cyanescens (Bull.) Quel. Gyroporus castaneus (Bull.) Quel. 

Gen. GYRODON Opat. 

Gyrodon lividus (Bull.) Sacc. 

Gen. BOLETINUS Kallbr. 

Boletinus cavipes (Opat.) Klotsch 


Phylloporus rhodoxanthus (Schweinitz) Bres. 


A. Edules Fr. 

Boletus edulis (Bull.) Fr. 
reticulatus (Schaeff.) Boud. 
pinicola Vitt. 

? aereus (Bull.) Fr. 

B. Rhodoporus Qu€L 

Boletus felleus (Bull.) Fr. 

C. Caerulescenti 

Boletus luridus (Schaeff.) Fr. 
erythropus Fr. non Pers. 
purpureus Fr. 

Satanas Lenz 

Queletii Schulz, var. lateritius Bres. & 
calopus Fr. 

Boletus albidus Roques 

appendiculatus (Schaeff.) Fr. 
regius Kxombh. 

? fragrans Vitt. 

{Ixocomus Quel.) 

{a) Annulati 
Boletus luteus (L.) Fr. 
elegans (Schum.) Fr. 
flavidus Fr. 
viscidus (L.) Fr. 
tridentinus Bres. 

{h) Exannulati 
Boletus granulatus (L.) Fr. 
bovinus (L.) Fr. 
variegatus (Swartz) Fr. 
sulphureus Fr. 
piperatus (L.) Fr. 

1 89 


Boletus aestivalis (Paul.) Fr. — pale form of B. appendiculatus, 
alutarius Bx,—B.fellea, 
aurantipori^^lows^^B, tridentinm. 
candicans Fr. — ^identity doubtful; probably B. albidus. 
carmsus Rostk.=j 5 . hadius. 
collinitus Fr. — probably B. luteus without ring. 
cruentusVGiit.=^B, impolitus, 

elegans Fr. — the valid name would appear to be Boletus Grevillei Klotsch, Linnaea, vii (1832). 

For the present the more familiar name may stand. 
erythropus Pers. non Fr.=R. Queletii. 

flavus (With.) Fr.— usually assumed to be identical with B. elegans. 
laricinus B. viscidus. 

nigrescens Roze & Rich.=R. crocipodius Let.; of the many epithets applied to this species by 
modern authors, each has some point which appears to invalidate its priority. B. crocipodius 
is based on a good figure by Letellier (1838). 
olivaceus (Schaeff.) Fr.=R. calopus. 

pachypus Fr. — identity doubtful; either B. calopus or B. albidus. 
paludosus Mass. — uncertain; may be a form of R. badius. 

pruinatus Bv. — a doubtful species midway between B. chrysenteron and B. versicolor, 
purpurascens Rostk. — looks like a form of B. versicolor, 
pusio Howse ex B, & Br. — not adequately described. 

Queletii Schulzer var. rubicundus Maire=var. lateritius. The type has not yet been recorded in 


radkans Yt.^^B. albidus. , 

rhodoxanthus (Krombh.) K.2LVLGDb.=^B. purpureus. 

Rostkovii Fr.— named by Fries from PI. 18 in Part iii of Sturm’s Die Pilze Deutschlands. Gilbert 
thinks it may be the same as B. tumidus Fr., but further observation required. 
rubiginosus Fr.— suggests form of B. reticulatus. ^ 

rugosus Fr.— Fries in error adopted Boletus rugosus from Sowerby, who under this name illustrated 
a Polyporus which looks like Daedalea biennis (Polyporus rufescens). In Hym. Fries refers to 
Sowerby, t. 420, but this plate is named Boletus lactifluus and is obviously B, granulatus. 

Fr.— uncertain what this refers to. 
sanguineus ('With.) Quel. = B. cramesinus Sect, 
sanguineus 'With, sensu versicolor. 

scaber Bull. — it is. an almost universal tradition that this refers to the common boletus of birch 
woods. Some French authors maintain that scaber should be adopted for a less common 
boletus described by Kallenbach ns pseudo-scaber. This epithet had been previously used and 
gives place to B. Carpini originally described by Roman Schulz as a vnriety of scaber. 
spadiceus — excluded from list to await clearer definition. It is probably only a form of sub- 
tomenfosus with white flesh and very prominent anastomosing ribs on the stem. 
spkaerocephalzis Bnrln. — a luxuriant form of R. 
tenuipes (Cke.) 'Mnss.= B. cramesinus. 
variecolorB. & Br. — uncertain what this refers to* 

vaccinus Fr. — from the description would appear to be a form of Gyroporus castaneus. 

Revised List of British Agarics and Boleti 

(Xerocomus Quel.) 

Boletus chrysenteron (Bull.) Fr. 
versicolor Rostk. 
subtomentosus (Schaeff.) Fr. 
parasiticus (Bull.) Fr. 
pulverulentus Opat. 
badius Fr* 
cramesinus Seer, 
impolitus Fr. 
rubinus W. G. Sm. 

Trachypus Bataiile 
(Krombholzia Karst.) 

Boletus scaber (Bull.) Krombh. 
var. coloripes Singer 
holopus Rostk. 
versipeliis Fr. 
duriusculus Schulzer 
crocipodius Letell. 

Carpini (R. Schulza) Pearson 

190 Transactions British Mycological Society 

Gyrodon caespitosus Mass. = caespitose form of Boletus pulverulentm, with tubes not fully developed. 
sistotrmaVx. = Gyrodon lividus, 

fuhellus McWeeney — ^probably Boletus versicolor with tubes not fully extended. 

Most of the British records of Gyrodon probably refer to young specimens of Boletus when the 
tubes are short and the pores more or less labyrinthine. G, lividus is the only fully authenti- 
cated species. 

Gyroporus lacteus {Lt6v.) Quel. = white form of G. cyanescens. 
fulvidus (Fr.) Pat. = pallid form of G. castaneus. 

Phaeoporus porphyrosporus (Fr.) Bat. — ^replaced by Porphyrellus porphyrosporus, the generic epithet 
Phaeoporus having been used by Schroeter for another genus of polypores, 

Ij'/opw^—replaced in Boletus^ the only species felleus having too close an affinity with the Edules 
section to be put into a separate genus in spite of its pinkish spores. 

{Accepted for publication i August 1946 ) 

[ I9I I 




Royal Botanic Gardens, Kew 

(With 5 Text-figures) 

Many of the names applied by Berkeley, Cooke and Massee to British 
Agarics have fallen into disuse because the published diagnoses were so 
inadequate that the species could not be recognized again with confidence. 
Preparation of a revised list of British Agaricaceae, undertaken in collabo- 
ration with Mr A. A. Pearson, necessitated reconsideration of these 
forgotten species and made it desirable to examine such material of them 
as was available, with a view to supplementing the original descriptions 
by whatever microscopic characters were still discernible. It is hoped that 
some of these dubious names will be reduced to synonymy and others 
revived as valid designations of well-defined species . Some of them probably 
refer to rare fungi that have not been collected by any competent mycologist 
since they were first described. This paper deals only with white-spored 
and pink-spored species. 


The material examined consists of dried specimens, mostly of type collections, 
from the herbaria of Berkeley, Cooke and Massee now preserved in the 
Gryptogamic herbarium at Kew, Wherever practicable, a small fragment 
of pileus was detached, soaked in lo % potash solution until softened and 
swollen to approximately fresh condition, transferred to dilute gum arabic 
solution and sectioned with the freezing microtome at right angles to the 
surface. The presence or absence of a specialized surface layer on the pileus 
was thus determined. Marginal portions of the gills were mounted in 
10 % potash solution and examined for cystidia and spores and, in white- 
spored species with abundant spores, a separate mount was made in 
Melzer’s reagent. A distinct dark blue coloration of the spore wall in this 
medium was observed only with Armillaria jasonis Cke. & Massee, Mycena 
Massee, zndi Pleurotus laurocerasi & Br.; spores of other species 
were unaffected or stained yellowish brown. Care is necessary in inter- 
preting observations made on such material, most of which has been dried 
for from fifty to one hundred years and is often in a fragmentary or poorly 
preserved condition. The spores observed have not been shed naturally 
and many of them must have been immature when dried, hence they may 
often not have attained their characteristic size and shape. Moreover, the 
gills are frequently found to bear as many as three totally different types 

192 Transactions British My cological Society 

of basidiospore, presumably because several collections had been carried 
in the same basket or vasculum and had shed spores on to one another. 
Many specimens also yield spores of various moulds, some probably 
parasitic, and here normal basidiospores can hardly be expected to be 
present. Hence one can only feel confident of identifying the right spore 
if one type clearly predominates over all others; only rarely can mature 
spores be found still attached to basidia. Cystidia, even of the thin-walled 
vesicular type, can usually be recovered in potash but in a few specimens 
the entire hymenium seems to have completely disorganized. Basidia are 
usually clearly discernible but the number of sterigmata is often doubtful. 

No specimens can be found at Kew of the following doubtful species: 
Lepiota emplastrum Cke. & Massee, Tricholoma duracinum Cke., Clitocybe 
zygophtla Cke. & Massee, C. pergamems Cke., C. occulta Cke., Collybia 
thelephora Cke. & Massee, Mycena consimilis Omphalia directa B. & Br., 
Hygrophorus aromaticus Berk., Entoloma liquescens Cke., E. fertile Berk., Molanea 
fulvo-strigosa B. & Br., Nolanea rubida Berk. 

(1) Lepiota polysticta Berk. 1836, p. 9. 

The type locality of L, polysticta was Cotterstock, Northamptonshire^ 
26 July 1828, and this collection cannot now be identified. Specimens, exist 
in Berkeley's herbarium labelled 'King's Cliffe’, and 'Coed Coch, 
Mrs Wynne Sep. 1865', and there is also an unlocalized collection now 
represented by two vertical sections of sporophores and two pressed speci- 
mens each consisting of a stipe and the skin of half a pileus about 3*5 cm. 
across. This sheet blears a pencil sketch of a spore with the dimension 
00025. It is labelled in Berkeley's hand- writing ^Ag. polystictus Berk.' and, 
if not type material, is at least a collection evidently studied in detail by 
the author of the species. This material was therefore chosen for re- 
examination. Kew also has two collections from the Hooker herbarium, 
labelled 'King's Cliffe ex herb. Berk.'; though authentic, these cannot be 
type material. 

The skin near the centre of the pileus consists of hymeniform cells 
measuring about 10 x 5^ resting on a tissue formed of narrow interwoven 
hyphae. Over the remainder of the surface the cuticle is broken up into 
rather conspicuous dark brown appressed squamules about 1-5 mm. across. 
The basidia are four-spored and the spores hyaline, non-amyloid, elliptical, 
6“8 X 3-4 /X, without a germ pore. The gill edge bears vesicular colourless 
cystidia approximately 25 x io-14/x. The scales on the stipe, mentioned in 
the diagnosis, are no longer apparent. 

This is quite a distinct fungus from that figured by Cooke as pL 41 (30) 
which has no scales on the cap. Berkeley’s species appears to fall in 
Kiihner's section Ovisporae oi Lepiota (Ktlhner, 1936) and may well be 
identical with i. clypeolarioides Rea. 

(2) Armillaria jasonis Cke. & Massee in Cooke, 1888. 

The type collection consists of a single cluster of five mature sporophores 
with a number of young ones growing out of a small mass of woody debris. 
The sheet is endorsed in pencil 'Carlisle Dr Carlisle figd.' and 

British Species of Agaricaceae. R. W. G. Dennis 193 

across this has been added in ink ^ Ag. {Armillaria) Jasonis C. & M, type’. 
The material is in good condition and sections of the pilens show it to be 
covered by a zone of rich brown globose or pyriform cells 1 5-2511 across and 
up to five cells deep. Below this and of the same colour is a region of 
similar but collapsed cells sharply differentiated from the underlying flesh 
which consists of rather stout colourless loosely interwoven hyphae about 
thick. Spores are abundant, hyaline, ovoid or slightly asymmetrical 
with distinctly amyloid walls 6-8X3-4/X. There are no cystidia. 

Fig, I. Armillaria jasonis. Section through surface layers of pileus, X 750. 

Armillaria jasonis is evidently Lepiota amianthina (Scop.) Fr., as already 
suggested by Boudier (1907)5 and by Quelet, Maire and Rea in Pearson 
(1935). It may be compared with var. Kiihner (1936), also 

found on rotting stumps, if that form is worth a varietal name. 

(3) Tricholoma circumtectum Ck.^^ & Massee, in Gooke, 1883— 9^5 P 3 ^ 2 * 
The type collection contains three dried and rather flattened sporophores 
labelled in Cooke’s handwriting - Ag, {Tricholoma)lLty^ Aug/88’, Attached 
to the same sheet is a copy of Cooke’s pL 1125 (1182) which bears the date 
Sept. 1888 and therefore presumably does not represent these specimens. 





OqO 0 c>on n 0^ 

Fig. 2 . A, Spores, cystidia from gill edge, and section of cuticle of pileus in Lepiota polysticta, 
B, Spores of Armillaria jasonts. G, Basidia and spores of Tricholoma circumtectum. D, Spores, 
basidium and section of surface of pileus in Tricholoma tenuiceps. E, Spores and section of 
surface layers of pileus in Clitocybe obscvrata. F, Spores and basidia of Laccaria nana. G, Spores, 
basidia and surface hyphae in Hygrophorus cerasinus. H, Spores, basidia and section of cuticle 
in Hygrophorus micaceus. Spores, basidia and section oi cnPiclQ m Hygrophorus ventricosus. 
K, Cystidia oi Collybiapsathyroides, L, Spores of Collybia eustegia, M., Spores and cystidium (?) 
of Collybia steverisoiiiu Ail >^ 750, 

British Species of Agancaceae. R. W. G. Dennis 195 

The material is in rather poor condition and had evidently begun to decay 
before being dried as the tissues are much contaminated with bacteria. It 
yields abundant non-amyloid hyaline broadly elliptical spores 5-7 x 2'5~4 /a 
borne on rather elongated basidia approximately 25-30 x 5-7 ja. Very few 
basidia still retain sterigmata but some at least appear to have been two- 
spored. There are no cystidia. The surface of the pileus consists of inter- 
woven hyphae with no distinct cellular cuticle. In Pearson (1935), Make 
suggested the fungus was T. atrosquamosum. Quelet thought it was T. argp'~ 
rdceum (Bull.) Fr. The former should have sterile dark cells on the gill edge, 
not seen in Cooke’s material; the latter identification may be correct but 
it is unwise to conclude more than that T. circumtectum represents a member 
of the T. myomyces group, probably the form called T*. scalpturatum (Fr.) 
Quel, by Bresadola (1881-92) and by Konrad and Maublanc (1924-37). 

(4) Tricholoma tenuiceps Cke. & Massee in Cooke, 1883-91, p. 398 (? 1891). 

Of this species there are preserved portions of six or seven sporophores, 
three still united at the base, labelled ""Ag, {Tricho,) tenuiceps C. & M. Kew 
July 1888’. The surface of the pileus consists of prostrate interwoven 
hyphae with brown contents, apparently coated with colourless gelatinous 
material which swells in potash. There is no sharp distinction between 
cuticle and flesh. The basidia are four-spored; the spores globose to sub- 
globose, non-amyloid, 7-8 x 6-7 fi. There are no cystidia. 

Evidently this is T. aggregatum (Schaeff.) Quel, as stated by Rea in 
Pearson (1935). Make suggested it might be a form of Collybia platyphylla 
(Pers.) Fr., influenced no doubt by the cord-like mycelial strands indicated 
on Cooke’s pi. 1121 (1166), but the globose spores are not those of that 

(5) Clitocybe obscurata Cke, 1909. 

The specimen of C. obscurata in Kew Herbarium consists of a single 
pileus, now 4 cm. across, with strongly decurrent gills. The lower part of 
the stipe is wanting and the pileus is firmly glued to the sheet by the whole 
upper surface. Hence the nature* of the cuticle could only be investigated 
near the margin, where it is composed of rather stout, closely interwoven 
hyphae, 3-4//, thick, forming a zone some 30/U, deep, overlying the rather 
spongy loosely woven flesh. The spores are hyaline, broadly elliptical 
non-amyloid, 5-6 (-8) x 3*5-4 (-5)^. This specimen is labelled 'Miss 
Graham coll. Clitocybe obscurata Cke. on the ground Edmond Castle, 
Carlisle, Oct, 1908’. 'Reed. 14. xi. 08.’ According to the diagnosis the 
type collection was made in September 1908 by the same lady in the same 
locality; possibly there has been some confusion of dates. As regards 
microscopic characters there seems nothing to separate this collection from 
C. clavipes (Pers.) Fr. to which Cooke thought it allied. His figure (Cooke, 
1909? t. 5 c) could represent an old specimen of that species in which the 
swollen base of the stipe was not developed. From C, subinvolutus W. G. Sm. 
it is distinguished by the absence on figure or specimen of any indication 
of an involute margin. 


196 Transactions British Mycological Society 

( 6 ) Laccafia nana ^ 

In view of Lange’s discovery (Lange, 1935-40) of a fungus closely 
resembling i. but differing from Massee’s diagnosis mainly in spore 
size and number of sterigmata, it seemed desirable to re-examine the type 
material of that species. The collection, labelled 'Laccaria nana sp. nov. 
K.G. 3/12 ’ consists of four dried sporophores in which the structures of 
trama and pileus are no longer distinguishable. The basidia are, however, 
well preserved, long and slender, about 10x3^, each with two well- 
developed sterigmata 6-7 ft long. The spores are spherical, 9-12^6 across, 
bearing spines 2 /x long, about thirty of which can be counted in the 
circumference of a spore 12/x in diameter. Lange’s fungus is thus clearly 
distinct from Massee’s. 

(7) Hygrophorus cerasiniis licYk. 1836, p. 12. 

Berkeley’s herbarium contains portions of at least two pilei, of diameters 
2*7 and 4*5 cm. respectively, with stipes as far as visible 4-8 x 0-4 and 
3*7 X 0*9 cm., glued by their lower surfaces to the same sheet as four 
vertical sections of sporophores with pilei 2-6, 4*4, 4*3 and 4*1 cm. across. 
All are riddled by mites and the gills of the three larger sections originally 
some 2~3 mm. broad, now consist mainly of frass. Those of the smallest 
section, though very narrow, are fairly well preserved. The sheet is 
labelled cerasinus Berk. Winkbourn, Notts. 1833’ and is evidently type 

The surface of the pileus consists of slender brown hyphae embedded in 
mucilage. The basidia were apparently four-spored but sterigmata are not 
well preserved. The spores are hyaline, broadly elliptical and strongly 
apiculate, 8-9 X 4-6/X. There seems no adequate grounds for attempting 
to separate this species from Hygrophorus {Limacium) agathosmus Fr. The 
name Agaricus agathosmus was coined by Fries (Fries, 1815, p. 17) for a 
fungus distinguished from Ag. pustulatus Persoon (Persoon, 1801) by its 
pleasant smell, colour and size. In the SystemUy however (Fries, 1821, i, 34), 
he changed his mind and cited A, agathosmus as a synonym of A, pustulatus 
Pers., making no reference to the odour. Later (Fries, 1836, p. 325) he 
changed his mind again and re-established Hygrophorus agathosmus as a 
distinct species. Secretan (1833) gave detailed descriptions of a fungus 
which he named, no doubt correctly, Agaricus agatJiosmus Hence 
Hygrophorus agathosmus (Fr. ex Secretan) Fr. is apparently the valid name 
for the species. 

(8) Hygrophorus micaceus B. & Br. Notices No. 1779, March 1879. 

The type sheet bears five poorly preserved sporophores andi traces of 
others and is labelled Hygrophorus micaceus & Br. Coed Coch. Oct. 1878’. 
The largest remaining pileus is 4 mm. across with a stipe 18 x 0-5 mm. and 
has a cuticle of brown, globose or broadly pear-shaped cells, approximately 
15-20/X across, which perhaps gave the "micaceous’ appearance to the 
surface. The basidia are slender, possibly only two-spored, with spores 
subglobose, 4-5 x 3‘5~4jtx. There are no cystidia. This seems to be a distinct 

British Species of Agaricaceae. R. W. G, Dennis 197 

species closely related to Omphalia atropuncta (Pers.) Quel, which Lange 
(1935-40) has transferred to 

(9) Hygrophoms ventricosus B. & Br. Notices No. 1777, March 1879. 

Berkeley’s herbarium contains two well-preserved sporophores under 
this name accompanied by two vertical sections, from one of which the 
pileus has been broken away and lost. All are firmly glued to the same 
sheet labelled ^Hygrophoms ventricosus B. & Br. Coed Coch 1878’. The 
existing pilei measure 2*6, 2-3 and 2-4 cm. in diameter and the stipes 
below the deeply decurrent narrow gills measure 5-o x 1*0, 4*4 x 1*2 and 
5*0 X 1*0 cm. respectively. There is no distinct cuticle, but the surface of 
the pileus consists of closely interwoven hyphae about 4-5 /T in diameter 
with no mucilaginous outer covering. The basidia seem to have been two- 
spored and the spores hyaline, non-amyloid, 7-10 x 4-5 /z, with ^ well- 
developed basal apiculus. The upper figures in Cooke’s pL 897 (901) no 
doubt represent the above collection and were thought by Make in 
Pearson (1935) to be H. virgineus. This seems likely enough for, according 
to Lange, two-spored forms of that species do exist. 

(10) Collybia eustygia Cht. 

The type packet contains one and a half dried sporophores, the larger 
now of maximum diameter 3*3 cm. with a stipe 8 mm. thick, and is 
labelled ^Ag. {Colly bia) eustygius C. & M. Whitfield 1890’. Pilei and stipes 
alike are dark chestnut brown shading to black, the gills dark grey to 
almost black. The surface of the pileus consists of interwoven hyphae with 
no distinct cuticle. The basidia are poorly preserved, mostly about 30 x 6 jx 
with sterigmata no longer visible and all have deep brown contents, giving 
the blackish hue to the gills. There are no cystidia. The spores are hyaline, 
non-amyloid, subglobose 5-6 X 4-5/^. 

From C, rancida Fr. this species is clearly distinguished by its much 
stouter stature and subglobose spores. C. eustygia is probably a slender 
form of the fungus described by Lange (1935-40) under the name Tricho- 
loma crassifolium (Berk.) Ricken and the lower central figure and upper left 
section in Cooke’s pi. 1146 (1185) accords reasonably well with Lange’s 
pi. 25 The fungus is, however, clearly not Ag, crossifolius Berk, as Lange 
admits. Colly bia apparently differs from Tricholoma immundum Berk, 

sensu Cooke, pi. 81 (61) and Bresadola (1881-92, pi. 156) only in its 
rooting base. Unfortunately, though there is abundant material of 
T. mmundurn in Berkeley’s herbarium labelled 'Moelfre uchaf, Denbigh- 
shire Oct. 1859’ it all appears to have been collected in bad condition 
with almost sterile hymenium bearing mainly the subglobose, spiny, 
non-apiculate spores of a parasite. Occasional subglobose smooth apiculate 
spores do occur and one apparently still attached to a basidium measures 
7 5*5/^* The hymenium oi T. immundum liz.s the same brown cell contents 

as Coilybia eustygia, Bresadola, and both Make and Rea in Pearson (1935) 
agree in regarding Tricholoma immundum as the same as Ag.fumosus Persoon. 
There is, however, some doubt which of the blackening species of Tricho- 
loma or Coilybia Persoon had, and also whether Fries used the name 

198 Transa^tiom British Mycolo^ 

Agarkus fumosus in the same sense as Persoon. It is proposed, therefore, 
to adhere in the meantime to the name Trickoloma immundum Berk. 

(ii) Agarkus cmssifolius i860. 

The type collection of this species was made at Winkbourn, Notts and 
has not been preserved. Berkeley’s herbarium contains a dried, pressed 
specimen consisting of the cuticle of a pileus now 7 cm. across, with stipe 
5x1-4 cm. tapering to the base, and a vertical section showing sinuate 
gills 7 mm. broad. These are labelled Cecil H. Spen 

Percival’ and are evidently the collection published in Notices 1503*, 
1876. There is also a water-colour drawing of three sporophores, the 
largest pileus 6*2 cm. across, labelled ^ A. pachyphyllus noW which is the 
source of Cooke’s pi. 93 (92) and presumably depicts the Winkbourn 
material. Both Spencer Percival’s fungus and the unlocalized drawing have 
a slightly scurfy fibrillose pileus like that of Trkholoma of the Myomyces 
group. This, taken in conjunction with the occurrence of the type in fir 
woods, is sufficient to disprove the suggestion by Quelet and Maire in 
Pearson (1935) that crassifolius is Trkholoma georgiL That it is not 

the fungus for which the epithet has been used by Bresadola and 

Lange is clear from Berkeley’s diagnosis, and from the specimen in his 
herbarium if that be accepted as authentic. Unfortunately, it has been 
much eaten by insects and its spores cannot be identified with certainty, 
though there are a few hyaline elliptical spores 6-7 x 4 ft present on the 
gills. The gills are now brown, not grey or black as in T immundum, and the 
surface of the pileus consists of matted hyphae about 3/x in diameter. 

{12) Collyhiaps 1883. 

The type collection, Epping Oct. 1880, contains the halves of a single 
sporophore with pileus now 1-4 cm. across and 1-0 cm. high, very thin 
flesh and broadly adnate gills 7 mm. deep at their attachment to the stipe. 
There remains about 1-5 cm. of a hollow stipe, some' 2 mm. thick, but the 
continuation of the stipe has been indicated on the sheet by a pencil outline 
to a total distance of 6-3 cm. from the pileus. The cuticle of the pileus is 
much disorganized but appears to have consisted of fine interwoven hyphae 
embedded in mucilage. The gills have an uneven surface and look abnormal. 
No spores can be found corresponding to those mentioned in the diagnoMs 
'white, elliptical 15 x7ft,’ there remain only the broadly elliptical or 
sudglobose hyaline spores of a parasite measuring 5-6 x 4-5 ft and brown 
spores I2“I6 X 8-iOft obviously alien, scattered rather freely over the gills. 
Gystidia are very abundant on the gill face and edge; most are pyriform 
and probably once bore a crown of spines as in many species of Mycena, 
for their apices are now obscurely punctate. The fungus was probably 
a diseased condition of some Mycena and the name should be discarded. 

(13) Colly bia stevensonii B. & Br. Notices No. 1497, Jan. 1875. 

Berkeley’s herbarium contains two sheets of this species each bearing 
two specimens, a vertical section and a half pileus, presumably of the same 
individual. All are much damaged by insects so that the gill attachment 

British Species of Agaricaceae. R. W. G. Dennis 199 

is no longer discernible. The large pileus is now 1*55 cm. across, the longest 
stipe 4*5 cm. x i mm. and the gills 3 mm. broad, but this may be an 
underestimate owing to the edge being broken away. The collection was 
labelled by Berkeley \Ag. {Collybia) Andersoni Glamis’ and bears also a 
separate label ‘described under the name of Agaricus stevensonii B. & Br.* 
Ann. Nat. Hist. No. 1439 M.C.G.’ Neither Berkeley and Broome, Cooke, 
pL 199B, Cooke (1883-91) nor Stevenson (1886) state the size of the spores; 
Rea’s (1922) measurement, lo-i i x 7-8 /x,, is copied from Massee (1893). 

The surface of the pileus was described as viscid and consists of slender 
hyphae, 2-3 p thick, embedded in mucilage; that of the stipe is smooth and 
free from cystidia-like hairs. Numerous vesicles, possibly to be interpreted 
as thin-walled cystidia are scattered over the gill face. They are elliptical 
in outline and some have brown oily contents. Spores are very scanty, the 
few found were hyaline, non-amyloid, and measured 6-8 x 3-4/^. The 
stature and long rooting base of this fungus suggest Marasmius esculentus 
(Wulf.) Karst, or M, conigenus (Pers.) Karst, but the viscid pileus is not 
that of those species. It seems likely that the type collection was abnormal 
or even diseased. 

(14) Mycena berkeleyi Massee, 1893, p. 104. 

The type consists of three specimens labelled by Berkeley ^Ag, {Mycena) 
excisus Hothorpe Nov. 15, 1881’ and a water-colour sketch by R. E. 
Berkeley which was faithfully reproduced as Cooke, pi. 224 (148). Spores 
are very abundant, broadly ellipical; 7-10 x 5-6 /x with distinctly amyloid 
walls. The basidia were probably four-spored and cystidia are no longer 
apparent but Maire’s suggestion in Pearson (1935) that M. berkeleyi was 
a form of M. galericulata (Scop.) Fr. appears highly probable. 

(15) Mycena paupercula Berk. 1836, p. 57. 

Berkeley’s herbarium contains two sheets under this name viz: 

{a) Agaricus pauperculus Clifton, Notts, eight specimens originally collected 
in excellent condition, sporing freely but now much riddled by insects. One 
pileus has been lost, the remainder now measure, 2, 2*5, 4*5, 3, 4, 4 and 
3 mm. in diameter respectively, with stipes 8, 4 ( + 7 mm. of root), 5 ( + 2), 

5 ( + 2), 8 ( + 3), 7 ( + 3):. 7 ( + 4). XI ( + 20) mm. long and 0-5-0-75 mm. 
thick. These are presumably the types, collected 20 Sept. 1832 and are 
accompanied by a water-colour sketch of seven pilei growing on decayed 
wood or detached to show the long rooting base. The sketch is labelled 
' A. pauperculus nob.’ and is the original of Cooke’s pL 231 (236). 

{b) No. 3158 Ag. pauperculus Berk. Texas C. Wright, seven specimens 
macroscopically similar to {a) and apparently growing on dead wood but 
without the long rooting base. Both in cuticle and spores they closely 
resemble the types but the pilei are much more distinctly striate. 

The following observations relate to collection (cz). The surface of the 
pilei consist of thin- walled, rather swollen, radiating hyphae containing 
dark brown granules. The basidia are four-spored clavate, about 20 x 5-6 ji6 
with non-amyloid spores 7-8 X 3 -5-5 /A. No cystidia have been detected. 
The surface of the stipe is smooth. 

200 Transactions British Mycological Society 

If this is a Mycena it would appear to fall in Kiihner’s (1938) subgenus 
Para-MycCTa, section Omphalariae, nudae. It differs, however, from any 
species there described in its narrow ascending gills, habitat and rooting 
base. Probably it is a good species which should be sought for and com- 
pared with Collybia muscigena (Schum.) Fr. 



Vig, 3. Omphalia abhorrens. Section of ptieus X175. Cuticle, b, Spongy tissue, c. Compact 
trama and subBymemuin. d, Hymeniuni, e, Cuticle in section x 750. Cuticle in surface 
view X750. 

(16) Omphalia abhorrensB. & Br. Notices 1751, March 1879. 

The type collection contains twelve sporophores, with stipes up to 
15x1-5 mm. and pilei up to 10 mm. across, labelled 'Ag. [Omphalia) 
abhorrens B. & Br. Coed Coch, odor stercorarius’. Microtome sections of 

British Species of Agaricaceae. R. W. G. Demis 20i 

the pileus show it to have a distinct yellowish brown cellular cuticle some 
15/X thick and 5-6 cells deep, covering a rather loosely woven, colourless, 
spongy flesh. The spores are subglobose, non-amyloid, 4-6 x 4-5 /x. There 
are no cystidia. This seems to be a well-defined species which should be 
easily recognizable when encountered again, but is it not the same as 
Phillips ? (Dec. 1878). 

(17) Omphalia alutacea Cke. & Massee in Massee, Dec. 1892. 

This species is represented in Cooke’s herbarium by four well-preserved 
sporophores of the type collection, labelled ^Agaricus {Omphalia) alutaceus 
C. & M. amongst grass Dinmore 1892’. The pilei are umbilicate with 
involute margins and now measure 10, 8, 8 and 6 mm. respectively with 
stipes 36, 34, 33 and 24x0*5-1 -5 mm. The gills are about i mm. broad 
and decurrent. There is no cellular cuticle, the surface of the pileus 
consists merely of more or less radiating interwoven hyphae. The basidia 
are four-spored; the spores hyaline, non-amyloid, broadly elliptical, with 
a basal apiculus 7-9 4 ‘ 5 “ 5 i^- There are no cystidia. The species, if 
distinct, is apparently close to Clitocype infundibuliformis (Schaeff.) Fr. 

(18) Omphalia telmatiaea Berk. & Cke. in Cooke, 1883-91, p. 93 and Cooke, 
March 1884, p. 66. 

The name was published without diagnosis but citing Cooke, pL 224 in 
Gremllea^ 1884. This species has been much confused. The diagnosis in 
Cooke’s Handbook cites ^ Cooke Ulus. t. 224 Ag. {Omphalia) affricatas B. & 
Br. in Handbook No. 216 not of Fries’. On referring to this we find cited 
Berkeley and Broome, Notices No. 994 and specimen from Aboyne, 
Aberdeenshire. Berkeley and Broome under No. 994 state ^Aboyne, 
Aberdeenshire, at the top of the Queen Hill, Aug. 9, 1862. Pileus... 
brown, then mouse coloured ... .’ 

Berkeley’s herbarium contains the following relevant material : 

{a) One sporophore 1*2 cm. across labelled ‘^Ag. {Omphalia) ajfricatus^ 
Aboyne 1862’, to which has been added, ‘Berk, non Fries’. This has sub- 
globose, non-amyloid spores 7-8 x ^-S/x, and no cystidia. 

{b) and {c) Two sporophores attached to labelled ^Ag. 

Fr. Glova, July, 1876, Dr Thomson’. 

{d) A water-colour drawing showing two views of a pileus 2 * i cm. across 
with stipe 15-20 mm. labelled ' Ag. affricatas pileo infundibuliform e um- 
bilicato hygrophano e fusco murino subtilita affricato. Lamellis distinctis 
distantibus abrupt! deliniatibus decurrentibus. Stipite compressis (?) basi 
tomentosis. Aboyne Aug. 9, Omphalia,^ Stipe and pileus alike are 

represented as slate-blue, somewhat as in Leptonia chalybaea. This is clearly 
a drawing of (<2) but bears no resemblance to Cooke’s pi. 256 (240) which 
figures specimens collected by Massee at Scarborough and not preserved 
at Kew. Berkeley’s drawing has been endorsed, apparently by Cooke 
=telmatiaeus 'Ecvk. & Cke.’ * 

Berkeley and Broome, No. 994, and Cooke, t. 256 (240), are surely not 
the same species and neither bears any resemblance to Fries’ (1867 t. 75) 
figure of Omphalia affricatas. The diagnosis of 0 . telmatiaea states ‘ Pileus 2 in. 

202 Transactions British Mycological Society 

broad, stem i^in. long’. This a^ees with Cooke’s plate but not with 
Berkeley and Broome, No. 994, which was described as f in. across. Hence 
the Scarborough collection must be accepted as the type of this species and 
no opinion on its identity can be based on the material at Kew. 

(19) Omphalia infumata B. & Br. Notices No. 1851, Feb. 1881. 

Berkeley’s herbarium contains three sporophores of this species, the 

largest pileus 8 mm. across, and one stipe from which the pileus has been 
lost, all attached to moss. The stipes measure 26, 23 and about 20 x i mm., 
swollen at the base to about 3 mm. The sheet is labelled 'A. [Omphalia) 
infumata B. & Br. Garthewin 1880’. There is no specialized cuticle, the 
surface of the pileus is formed of prostrate hyphae with slightly swollen 
tips; the stipe is smooth. There are no cystidia. The basidia are cylindrical, 
about 35 X 6ju., and probably four-spored though sterigmata are no longer 
clearly visible. The spores are broadly elliptical to elliptic-oblong or 
slightly inequilateral with a basal apiculus, non-amyloid, 8-10 x 4-5/4. 

(20) Pleurotus hobsoni Berk, i860, pp. 138-9. 

The type sheet bears eight sporophores attached to moss and labelled 
‘Agaricus [Pleurotus) B. Apethorpe, Sept. 1859’. The largest measures 

7 mm. across by 4 mm. from back to front; all are sessile with well- 
developed giUs now about 0-5 mm. wide. The surface of the pileus is 
formed of rather loosely woven hyphae. The basidia were probably 
four-spored with hyaline, non-amyloid, elliptical spores 6-10x5-6/1, and 
there are no cystidia. Unfortrmately the gills yield a rather mixed collec- 
tion of spores, but many of those present are certainly longitudinally 
ridged and this may well be Clitopilus pleurotelloides (Kuhner) Josserand. 
Berkeley apparently did not take a spore print so one cannot tell whether 
the spores were pink in the mass or not. In addition to the microscopic 
characters there is fair agreement in the macroscopic appearance of 
Pleurotus hobsoni and Clitopilus pleurotelloides as shown in the following 
comparative table : 

after Berkeley’s 

diagnosis Clitopilus pleurotelloides ^JCCox 6 hlg to Josstmiid 

1. Pileus membranaceous i. Pileus non-hygrophanous, dry 

2. Reniform or dimidiate 2. ± irregular, reniform, auricular 

3. Stemless 3* Soon becoming laterally attached or even re- 


4. Pale grey 4. Snow white 

5. Minutely downy 5. Finely pubescent, when yoimg, surface felted 

when old 

6. Gills rather distant 6. Gills fairly crowded 

7. Gills pallid 7. ^ills ivory white, then cream and pale ochraceous 

cream, a little tinged rosy 

8 . On larch stumps 8. On rotting or mossy bark 

9. Pileus i-~4 lines across ^ V 9. Pileus 4-15 mm. 

10. Margin involute 10. Edge subobtuse when young 

11. September ii. June-October.. 

In view of the poor condition of Berkeley’s collection it seems undesirable, 
however, to make a new combination for it in Clitopilus, It would seem 
wise to discard the name hobsoni Berk. 

20 $ 

British Species of Agaricaceae, R. W. G, Dennis 

(21) PleuTotus laurocerasi B. & Br. Notices No. 1854, 1881. 

The type consists of a single flattened, unstalked fructification about 
2 cm. across and 1-4 cm. from base to margin. Berkeley's label reads "Ag. 
(Pleurotus) laurocerasi B. on Laurel Coed Coch’ and to this Dr A. Pilat has 
added 'Pleurotus tremens Quel probabiliter identicus 29. xi. 1934'. P. tero- 
cerasi has no cystidia and amyloid, broadly elliptical smooth spores 
9-12 (”-13) X 7-8 ft. According to Quelet (1877) P- tremens had spherical 
aculeate spores measuring 6-7 ft and he so figured them. These recall those 
of P. palmatus (Bull.) Fr. as already pointed out by Lloyd (1901). 

Pilat (1935) says of this specimen "^En tant qu'on peut juger de ce 
mauvais exemplaire, c'est un champignon, dont le chapeau et les lamelles 
ont une consistence gelatineuse. Basides 20-28 x 5-6 ft. Sans cystides. 
Hyphes de la trame des lamelles a parois minces, gelatineuses, hyaline, 
densement et parallelement entrelacees, epaisses de 7-1 1 ft. Hyphes du 
chapeau analogues. Spores largement et subspheriquement ovoides, avec 
un apicule minuscule a la base, hyalines, 8-8*5 x 6-6*5 ft, des granules 

gdatineux colles a la surface, ce qui, leur donne une apparence squamu- 
leuse, Apparement une bonne espece, assurement fort rare.' 

There is a large number of subspherical spores of the size quoted by 
Pilat but as there are also many broadly elliptical large ones I have taken 
the former to be immature. The squamulose or wrinkled surface is not 
easily seen even in Melzer's reagent and cannot be detected at all in many 
of the larger spores. 

(22) Pleurotus ruthdeP^. & Br. Notices No. 1754, March 1879. 

Berkeley's herbarium contains two specimens under this name. The 
smaller is well preserved and labelled ' Ag, [Pleurotus) ruthae B. & Br. Coed 
Coch Oct. 1878 but there is also a larger, very much decayed, unlocalized 
fructification and a water-colour sketch dated ' Coed Coch '79 '. Dr Pilat has 
attached a note to each specimen determining it as Pleurotus petaloides (Bull.) 
Fr. The smaller sporophore has abundant, fusiform, thick-walled cystidia 
measuring about 80-100 x 12- 16 ft scattered over the gill and smooth, 
hyaline, non-amyloid spores 7-9 (-10) x4-5*5ft. It is no doubt P, peta- 
loides. Pilat (1935) accepts the smaller specimen as typical of that species 
and the large as approaching the var. geogonius (DC.) Pilat. 

(23) B. & Br. 1871, p. 5 

Pluteus spilopus was described by Berkeley and Broome from a dried 
specimen and figure sent them from Ceylon. Subsequently (Notices 
No. 1856, 1881), they published it from Britain. Berkeley’s herbarium 
contains a single sheet labelled 'Ag. [Pluteus] spilopus B. & Br.' bearing a 
pressed half sporophore and four vertical sections. This is unlocalized but 
can be identified with the Ceylon material as it has the No. 1167 quoted in 
the diagnosis. As the British collection is not available for comparison no 
final opinion can be expressed regarding its identification with P. 

As tropical fungi have seldom been found to be identical with temperate 


Fig. 4. A, Spores of Tncholoma crassifolm. and basidium. of Mycem berkeleyu f 

Gj Spores of Omphalia abhorrms. Spores of Omphalia alutacea. E, Spores, basidia and 
surface hyphae from pileus of Mycem paupercula. F, Spores and surface hyphae in Omphalia 
infumaia. G, Spores of Omphalia telrmiiaea (Aboyne specimen) . H, Spores, basidium and * 

surface section of pileus in Pleurotus hobsonL J, Spores of Plmrotus lamocerasi. K, Spores 
ofEntoloma bulbigenum (Sibbertoft collection). L, Spores, cystidium and cuticular cells of 
pileus of Entoloma thomoni. M, Spores and cystidia of Pleurotus ruthae. N, Spores, cuticular 
cells and hairs from pileus of Entoloma uymiae, O, Spores of Clitopilus sarnicus. P, Spores 
of All X750. ^ ^ ^ T 


British Species of Agaricaceae. R. W. G. Dennis 205 

species, however, it seems highly improbable that they were the same and 
P. should be deleted from the British list until confirmed by a fresh 


(24) Entoloma hulbigenum B. & Br. Notices No. 1937, March 1882. 

There has been deplorable confusion over this name. Berkeley and 
Broome in coining it gave no diagnosis but stated A, {Entoloma) bulbigerms 
B. & Br.; - 4 . Persoonianus PhilL GarL Chron, 1881, p. 874; A. sericeus Pers. Ic. 
et Descr. t. 6, f. 2. East Dereham, Rev. J. M. Du Port. Sibbertoft, Norths. 
Feb. 1882, Miss Ruth Berkeley." Just the plant of Persoon, who indicates 
in his figure the sclerotioid bodies at the base, though he does not mention 
them in the text. The name of A. persoonianus requires to be changed as 
there is a species of similar name. 'Fr. Hym. Eur, p. 25. Spores 0-0004 
0-0005 inch in diameter." The reference to Fries (1874) is to Ag. (Amanita) 
persoonii Fr. and seems scarcely to necessitate a change of name. Evidently 
Berkeley believed himself to be renaming Phillip’s fungus. His herbarium 
contains under this name only two dried sporophores and a vertical section 
from the Sibbertoft collection, accompanied by a water-colour drawing. 
The ‘sclerotioid bodies’ are merely enlarged bases of the stipes, which are 
shown to be distinctly floccose in the lower third. A fragment of a dead 
tree leaf is attached to the base of one stipe. This fungus has broadly 
elliptic-oblong, hyaline spores 11-12 x and broadly adnate gills 

tinted pink in the drawing. There are no cystidia. Probably it was a 
Marasmius or a Colly bia and is certainly not the same as the fungus figured 
by Phillips (1881) as having bottle-shaped encrusted cystidia and star- 
shaped spores suggestive of a form of Inocybe fibrosa (Sow.) Fr. Phillips 
identified his fungus with Agaricus sericeus Persoon and merely quoted 
Persoon’s diagnosis with the additional and evidently erroneous statement 
that the plant was an Entoloma, He gave no spore measurements. Phillips 
and Plowright (1881) however, in a contemporaneous note, stated that 
‘the bulb in our plant is much more decided than in Persoon’s figure 
•having a distinctly abrupt margin, and the pileus is not so fleshy. The large 
angular spores are as wide as the basidia and the hymenium abounds with 
large cystidia. Spores, including the angles; 0-015 mm.’ They still believed 
themselves to be merely renaming Persoon’s Ag, sericeus which Fries had 
placed as a variety of Ag, sericellus Fr. In view of the discrepancies noted 
by them, their meagre description and published figure may perhaps be 
taken as the diagnosis of a new species of which Du Port’s Norfolk collection 
will then be the type. 

It is noteworthy that in the same issue of Gremllea, Cooke (1881) 
published the name (Entoloma) persoonii Du Port, again citing both 
Ag, sericeus Persoon and the Norfolk fungus. His pi. 324 (315) is in the 
upper part a copy of Berkeley’s water-colour of the Sibbertoft collection, in 
the lower half a copy of Persoon’s figure of A, sericeus (Persoon, 1798, t. vi. 
fig. 2) as regards the section and the fructification to the left. That in the 
lower right corner is probably merely a product of Cooke’s imagination 
supplied to fill the space and balance the plate. Whether Cooke ever saw 
Du Port’s fungus is doubtful but the copy of pL 324 (315) in his collection 

2 o 6 Transactions British My cological Society 

of drawings at Kew bears a manuscript note to the effect that the spores 
measured 13-15 X 1 0-12 ft. 

Entoloma bulbigenum should be discarded as a name which originated in 

(25) Entoloma thomsoni B, & Br. Notices No. 1523, Feb. 1876. 

The type collection consists of four pressed sporophores mounted on two 
small sheets, each labelled "Ag, [Entoloma) Tliomsom B. & Br. West Far- 
leigh ’. The largest pileus has been much deformed in pressing, the others 
measure 2-7, 2*5 and 2-0 cm. and the longest stipe is about 4 cm. x 2 mm. 

The surface of the pileus bears vesicular cells 20-25 x about 15 ft which 
presumably gave it the 'velvety’ appearance. The spores are 7“8 (-9) x 4-6fi-5 
broadly elliptical and yellowish under the microscope. The hymenial 
elements are poorly preserved but there seem to be large numbers of 
thin-walled, pointed, vesicular cystidia. Clearly this is not an Entoloma or 
2l Rhodophyllus of any kind. The spores and free gills point to a Pluteus^ 
almost certainly P. cinereus Quelet (1884) and the name should therefore be 
Pluteus thomsoni (Berk.) comb. nov. 

(26) Entoloma wynniae B. & Br. Notices No. 1342, May 1873. 

Seven specimens from the type collection of E, wynniae are preserved in 
Berkeley’s herbarium, labelled [Entoloma) wynnei'B, & Br. Coed Coch 
1872 near A, costatus\ The largest pileus now measures 2*5 cm. in diameter, 
the longest stipe about 3*5 cm. x 3 m. The surface of the pileus is formed of 
vesicular pyriform cells interspersed with elongated swollen brown hairs 
up to 100x12 ft. The spores are of the Rhodophyllus tYpt 13-17^1 long, 
including the prominent apiculus and 6-8 ft broad. No cystidia were seen. 

[ 9 ,j) Molanea babingfonii B. & Br. Notices 680, May-June 1854. 

The type material labelled ^ A. Babingtonii Blox. Ag. [Molanea) Bloxami 
Berk. Rev. A. Bloxam Twycross Nov. 21, 1851’ is riddled by mites and 
consists of little more than a mass of frass. No Rhodophyllus spores were found 
and some, perhaps all, of the hair-like bodies on the cap are the brown 
geniculate conidiophores of a Dematiaceous mould. The Colly weston 
collection mentioned in Notices 903* is evidently that labelled ^Ag. 
Babingtonii^. King’s Cliffe, Oct. 2, i860’. It contains two well-preserved 
sporophores with abundant Rhodophyllus spores 16-20 x 8-1 o/x. The surface 
of the pileus is composed of inflated parallel hyphae running out into 
rather short swollen hairs with cells some 25 /x broad. Similar but longer 
hairs occur on the stipe. Mr Pearson thinks this is Nolanea fumosella 
(Winter) Sacc. [ = M. strigosissima 'R.t^i). 

(28) Massee 1897, p. 22. 

The type collection contains a single specimen with pileus 2*3 cm. across, 
conspicuously striate to the disk, with stipe 1-4 cm. x 2 mm. It is pressed 
flat but the gills were apparently adnate or adnato-decurrent. There is no 
specialized cuticle and there are no cystidia. The spores are of the Rhodo- 
type, 10-12 x 6-9/x. It is apparently a fovm oi Clitopilus cancrinus Pr. 


British Species of Agaricaceae. R, W. G. Demis 

(29) Clitopilus stilbocephalus B. & Br. Notices No. 1758, March 1879. 

The type collection contains two pressed sporophpres with pilei 2-5 and 
2-2 cm. diameter and stipes 4-6 cm. x i mm. and about 7-0 cm. x 1-5 mm. 
respectively, and a vertical section showing adnate gills 3 mm. broad. 
There are also pen and ink sketches of a campanulate pileus and of three 
sections, one showing broad, deeply sinuate gills, the others adnate ones. 

Fig. 5. Molanea babingtoniu A, Hair of stipe. B, Hairs of pileus. C, Surface cells 
of pileus, all X500. D, Spores x 750. 

On the same sheet is a manuscript Latin diagnosis similar to that published 
with in addition 'odore non ingrati’ and the note ‘sparkling as in d. ato- 
matus^: Cooke’s pi. 349 (324) lower figures was concocted largely from 
these sketches. The surface of the pileus seems to be formed of radiating 
hyphae with nothing to suggest the reason for the atomate appearance. 
The spores are of the Rhodophyllus type, 11-13 x 6“8jit. Possibly 
Entoloma griseocyaneum (Fr.) Quel. 

2 o 8 Transactions British Mycological Society 


Re-examination of authentic material of twenty-nine little known British 
Agarics lead to the following conclusions : 

(i) Lepiota polysticta Berk, probably equals X. clypeolarioides Rea. 

{9) Armillaria jasonis Cke. & Massee is Lepiota amianthina (Scop.) Fr. 

(3) T richoloma circumtectum Cke. & Massee is probably T scalpturatum 

Fr. sensu Bresadola, 

(4) T. temiceps Cke. & Massee is T. aggregatum (Schaeff.) Quel. 

(5) Clitocybe obscurata Cke, is probably a form of C. clavipes (Pers.) Fr. 

(6) Laccaria nana Massee is a distinct species correctly described by 


(7) Hygrophorus cerasinus Berk, is XX agathosmus Fr. 

(8) XX micaceus B. & Br. is a distinct species near Omphalia atropuncta 

(Pers.) Quel. 

(9) Hygrophorus ventricosus B. & Br. is a form of Xf. virginem (Wulf.) Fr. 

(10) Collybia eustygia Cke. is a rooting form of Tricholoma immundum Berk. 

(11) Agaricus crassifolius Berk, is probably a member of the Tricholoma 

myomyces group. 

(12) Collybia psathyroides Cke. was based on diseased material and should 

be discarded. 

(13) C. stevensonii B. & Br. was probably founded on an abnormal plant 

and cannot at present be recognized. 

(14) Mycena berkleyi Massee was probably M, galericulata (Scop.) Fr. 

(15) M. paupercula Berk, is probably a good species. 

(16) Omphalia abhorrens B. & Br. is probably Hygrophorus foetens Phillips, 

(17) Omphalia alutacea Cke. & Massee may be distinct. 

(18) 0 . telmatiaea Berk. & Cke.-must remain in doubt until the type is 


(19) 0 . B. & Br. may be distinct. 

(20) Pleurotus hobsoni Berk, may probably be the same as Clitopilus 

(Kuhner) Josserand. 

(21) Pleurotus laurocerasi B. & Br. is a distinct species. 

(22) P. ruthae B. & Br. is Pleurotus petaloides (Bull.) Fr, 

(23} Pluteus spilopus B. & Br. should be deleted from the British list of 
Agaricaceae. It is a good species from Ceylon. 

{24^ Entdoma bulbigenum Berk, originated in confusion. Agaricus per- 
sooriiarms PhlAi. probably was Inocybe fibrosa (Sow^ Fr. 

(25) Entoloma thomsoni B. & Br. is a Pluteus^ P. thomsoni (B. & Br.) comb. 

nov., identical with P. cinereus Quel. 

(26) Entoloma wynniae & Br. may be a good species. 

(27) Nolanea babingtonii Berk, is probably the valid name for M.fumosella 

(Winter) Sacc. 

(28) Clitopilus sarnicus Massee is probably a form of C. cancrinus Fr. 

(29) C. stilbocephalusB. 81 BT. m3.Y he Entoloma griseo-cyaneum (Fr.) Quel. 

My thanks are due to Miss E. M. Wakefield for criticism and advice, 
especially on questions of nomenclature, and to Mr A. A. Pearson for 
several suggestions regarding the identity of the more difficult species. 

British Species of Agaricaceae. R. W. G. Dennis 



Berkeley, M. J. (1836). Fungi in Hooker, W. J. Cryptogamia. The English Flora of Sir 
James Edward Smithy V, pi, 2. 

Berkeley, M. J. (i860). Outlines of British Fungology. London. 

Berkeley, M. J. & Broome, C. E. (1837-85). Moikes of British Fungi 
Berkeley, M.J. & Broome, C. E. (1871), The fungi of Ceylon. J, Linn, Soc.Lond, (BoL), 
XI, 495-567. 

Boupier, E. (1 907) . Quelques rectifications e t observations critiques sur les ‘ Illustrations 
of British Fungi’ de Cooke. Trans. Brit. myc. Soc. n, 150-7. 

Bresadola, J. (1881-92). Fungi Tridentini noviy vel nondum delineati, descripti, et iconibus 
illustratiy 2 Yols. Tvtnt. 

Cooke, M. C. (1881-91). Illustrations of British Fungi 

Cooke, M. C. (Dec. 1881). New British Fungi. Grevilleay x (no. 54), 4I-52. 

Cooke, M. C. (June 1883). New British Fungi. Grevilleay ' kj, 155. 

Cooke, M. C. (1883-91). Handbook of British Fungiy 2nd td. London. 

Cooke, M. C. ( March 1884). New British Fungi. xii, 65-70. 

Cooke, M. C. (March 1888). New British Fungi. Grevillea, xviy 77. 

Cooke, M. C. (Dec. 1890). New British Fungi. Grevilleay isiKy 40-2. 

Cooke, M. C. (1909). Fungus notes for 1908. Trans. Brit. myc. Soc. in, 109-10. 

Fries, E. (1815). Observationes Mycologicae praecipm ad illustrandam Flora Sueciam. 

Fries, E. (1821). Systema rnycologicumy i. 

FmESy E. Epicrisis Systematis Mycologici. Uppsala. 

Fries, E. (1867). Fones selectae Hymenomycetum nondum delineatorum. 

Fries, E. (1874). Hymenomycetae Europael 

JossERAND, M. (1941). fitudes sur les espdces frangaises du genre Clitopilus. Bull Soc. 
Linn. LyoUy x, 91-4, 104-12. 

Konrad, P. & Maublang, A. (1924-37). leones selectae Fungorum. Paris. 

KteNER, R. (1936). Recherches sur le genre Lepiota, Bull Soc. Myc. Franccy lii, I77-240, 
KiJhner, R. (1938). Le genre Mycena. Paris. 

Lange, J. E. (1935-40). Flora Agaricina Danica. 5 vols. Copenhagen. 

Lloyd, G. G. (1901). Pleurotus subpalmatus. Mycological Notes yViy ^1-2. 

Massee, G. (1892). New or Critical British fungi. xxi, 40-1. 

Massee, G. (1893). British Fungus Floray niy London. 

Massee, G. (1897). British Mycology. Trans. Brit. myc. Soc. i, 20-4. 

Massee, G. (1913). Additions to the wild fauna and flora of the Royal Botanic Gardens, 
Kew, XIV. Kew Bulletiny ^ 

Pearson, A. A, (1935)* Cooke’s Illustrations of British Fungi. Trans. Brit. myc. Soc. xx 


Persoon, D. C. (1798). leones et Descriptiones fungorum minus cognitorumy Fasc. 1 . 

Persoon, D. C. (1801). Synopsis methodka fungorum. Gottingen. 

Phillips, W. (Dec. 1878). Hygrophorus foetens nov.spec, Grevilleay vii, 74-6. 

Phillips, W. (Dec. 1881). Notes on some recent forays among funguses. The Gardeners' 
Chronicle (N.S.), xvi, 784. 

Phillips, W. & Plowright (Dec. 1881). New and rare British fungi. Grevilleay x 
(no. 54), 65-74. 

PiLAT, A. (1935). Atlas des Champignons de VEuropCy Tome n, Pleurotus Fries, Praha. 
QuiLET, L. (1877). Qnelques esp^ces de champignons nouvellement observees dans le 
Jura, dans les Vosges et aux environs de Paris. Bull. Soc. Bot. FrancCy xxrv, 317-32. 
Quelet, L. (1884). In Ann. des Sciences nat. de Bordeaux. Qpoted in Froc. Association 
Frangaise pour Lavancement des sciencesy Congris de Grenobky 
Rea, G. (1922). British Basidiomycetae. Cambridge. 

Secretan, L. (1833). Mycographie Suisse, n. Geneva. 

Stevenson, R. J. (1886). British Fungi {Hymenomycetes). Edinburgh and London. 
Winter, G. (1884). PilzCy Dr N. Rabenhorsf s Kryptogamen Flora von Deutschland, Oesterrekh 
und der Schweiz, h Nbth J. 

{Accepted for publication February 


[ 210 ] 



The Wellcome Physiological Research Laboratories^ Beckenham^ Kent 
(With Plates XII and XIII and 6 Text-figures) 


In these laboratories, the best medium for the growth of Streptomyces griseus 
and for streptomycin production by this organism has been found to be 
one containing mineral salts, glucose, peptone, and meat extract, described 
by Ainsworth, Brown, Marsden, Smith and Spilsbury (1947) as ^Belmont 
medium’. This paper is an attempt to outline the nutritional requirements 
of the organism and demonstrates that the success of Belmont medium is 
largely due to its mineral content and the added meat extract. 


All the experiments described below were performed at 28^^ C., the 
optimum temperature for growth as determined in these laboratories, and 
the strain used was the one employed for large-scale streptomycin produc- 
tion by Ainsworth al. (1947), National Collection of Type Cultures 
no. 71 87. The pH of all media was adjusted to 6-9 ± o* i before sterilization. 
Assay was carried out by the method of Brown and Young (1947), using 
Escherichia coli as the test organism, the results being expressed in micro- 
grams of pure streptomycin base. 

Mycelial dry weights were obtained either by collecting the mycelium 
and drying to constant weight in an oven at 100° C. or by freeze-drying 
the washed mycelia in tared aluminium receptacles. 

It will be convenient to discuss the experiments in three groups; the 
effect of metals upon growth, the nitrogen source, and growth-factor 

The effect of metals upon growth 

Belmont medium as used for streptomycin production normally contains 
added iron and copper as sulphates in addition to sodium nitrate, potas- 
sium dihydrogen pihosphate and magnesium sulphate. The inclusion of 
these substances being purely empirical it was felt desirable to investigate 
their several effects on the growth of the organism. 

In order to obtain some basis upon which to work, a sample of dried 
material obtained from mycelial mats grown upon Belmont medium was 
submitted to spectrographic analysis by the arc method. The principal 

Nutritional retirements of Streptomyces griseiis. Spilsbury 211 

elements detected were calcium, magnesium, sodium, silicon, iron, potas- 
sium and phosphorus, arranged in decreasing order of quantity as estimated 
from the intensities of the lines. A small quantity of copper was detected 
and also traces of aluminium, barium, strontium, lithium, rubidium, 
manganese and lead. The presence of zinc was doubtful. 

In addition to this analysis, a further preliminary investigation was 
undertaken by incorporating various elements at 0*4, 2*0 and lO'O p.p.m. 
concentrations in a synthetic medium containing ammonium nitrate as 
the nitrogen source at a level of i mg./ml. nitrogen together with the salts 
normally present in Belmont me&um. The medium also contained 
I % glucose and 50 mg./ml. tryptophan. It was found that the ions of 
lead, tin, uranium, vanadium, cerium, strontium, chlorine, iodine and 
fluorine had no appreciable effect at any concentration employed. Slight 
stimulation of growth appeared to result from ions of bromine at 2 p*p.m., 
zirconium at 10 p.p.m., and all concentrations of molybdenum employed, 
which latter result was independently confirmed by Miss E. Brookfield in 
these laboratories. Bismuth and lithium were slightly toxic at 2 p.p.m., 
aluminium, cobalt and nickel were toxic at 10 p.p.m. (also confirmed by 
Miss Brookfield), while cadmium proved to be extremely toxic at all 
concentrations employed. From the above data it was considered that the 
metals most likely to prove worthy of investigation were copper, iron, 
manganese, zinc and molybdenum, together with the major salt con- 
stituents. It is noteworthy that copper, manganese and zinc are recognized 
to be important in penicillin production by certain strains of Penicillium 
notatum and P. chrysogenum, that manganese has been shown by Robbins 
and Hervey (1944) to be essential to the growth oi Pjthiomorpka gonapo- 
dyides^ and that molybdenum has been shown by the author (unpublished 
data) to affect the sporulation of Penicillium notatum^ strain N.R.R.L. 
1249 B21, when added to Moyer’s glycerol molasses medium. There are 
also numerous references in the literature to the effect of these metals on 
the nutrition of fungi (see Stiles, 1946), but none as far as could be ascer- 
tained with respect to actinomycetes. 

The effect of copper ^ iron, manganese and zinc in Belmont medium 

The first experiment consisted of adding copper, iron, manganese and 
zinc as ^^Analar’ hydrated sulphates to Belmont medium from which the 
copper and iron had been omitted. They were added singly at concentrar 
tions of 50 and 10 p.p.m. of metal and in all possible combinations at 
10 p.p.m., a control being left without any trace metal addition. The media 
were distributed in 200 ml. quantities in pint milk bottles, the necks of 
which were closed with aluminium caps as described by Ainsworth al. 
(1947), The bottles were sterilized, inoculated from a suspension of 
Streptomyces griseus grown upon normal Belmont medium, and sloped. 
From the commencement of growth there were noticeable differences in 
the appearance of the growths resulting from different treatments. On the 
third day after inoculation the control bottle was completely covered by 
a whitish growth. 50 p.p.m. copper resulted in a thicker growth with a 

■■■ , 14-2 

212 Transactions British Mycological Society 

tendency to greyish appearance. 50 p.p.m. of zinc were inhibitory, the 
surface growth consisting of small scattered brownish coloni^, while much 
of the growth consisted of sphaeroidal submerged colonies similar to those 
normally obtained in shaken cultures in conical flasks. The effects of 
manganese and iron were similar to that of copper, except that in the case 
of iron the growth was definitely brownish. Concentrations of ro p.p.m. 
copper, manganese and iron resulted in growth of a heavier nature than 
the control or with the same metals at 50 p.p.m. Zinc at 10 p.p.m. did not 
cause such abnormal growth as at the higher concentration, but the 
appearance of the mycelium was whitish and granular. Mixtures of the 
metals tended to give an intermediate appearance; thus mixtures con- 
taining zinc possessed a tendency to granular growth, those containing 
iron to become brown, and those containing copper or manganese to 
become thicker, wrinkled and grey. It may be noted that the brown 
coloration obtained in this experiment was identical in appearance with 
that obtained by incubation at a temperature in excess of the optimum 
(Ainsworth et al. 1947) and was due to the waterlogging of the mycelium 
with the medium and not to any morphological change in the hyphae. 
This phenomenon appears to be intimately connected with the metabolism 
of the mould ; its reversibility with regard to temperature has been demon- 
strated, and it seems likely that a change in the constituents of the medium 
alters the optimum temperature for the production of aerial mycelium. 
The grey coloration obtained in this experiment resulted from free 
sporulation of the organism which might or might not precede the forma- 
tion of a brown mycelial mat. The white, granular appearance is typical of 
the vegetative phase of the organism on semi-solid media such as that of 
LePage and Campbell (1946). Good growth obtained in the control of 
this experiment cannot be interpreted as an indication that these trace 
metals are inessential to growth, since such elements would undoubtedly 
be added in the peptone and meat extract, and, indeed, the only interpreta- 
tion possible is that of the gross effect of further additions of trace metals 
added to a medium already containing a sufficiency of them to permit 
adequate growth. The spectrographic analysis of the organism confirms 
that copper and manganese are absorbed from the media and therefore 
probably possess some physiological function; the submerged growth 
resulting from zinc treatment indicates that this metal at least has a marked 
effect on the general physiology of the actinomycete, and the data obtained 
show that the trace metals present have a definite effect on both the 
amount of mycelium obtained and the titre of streptomycin produced 
therefirom. The titre, pH and mycelial dry weights of the cultures are 
presented, together with their analysis of variance in Table i, and the 
effects of the trace metals on titre and mycelial dry weight are represented 
diagrammatically after the method of Richards (1941) in Text-figs, i 
and 2. 

The general conclusions to be drawn from the data are that the presence 
of copper definitely increases the titre, the optimum concentration in the 
Belmont medium lying between 10 and 15 p.p.m. of added metal. Man- 
ganese has a slighter but still significant effect, and zinc decreases the titre. 

Nutritional requirements of Streptomyces griseus. Spikbury 21 3 

The best combiaation for streptomycin production appears to be copper- 
manganese mixture. Copper definitely antagonizes the bad effect of zinc. 
The combinations of manganese with iron and of zinc with iron would be 
expected to be bad, judging from their main effects, and fulfil expectation. 
Data of a similar nature were obtained after 12 days’ growth, and although 
not complete, they substantiate the above findings. lu general, it was 
observed in this experiment that factors tending to good titre also gave the 
highest pH values. This experiment was repeated with similar results. 

Table i. Effect of trace metals on growth, titre and pH. The data presented 
are the result of bulking three bottles from each treatment 

7 day 12 day 



Dry wt, t 

Probability ^g./ml. 




pH ftg./ml. 


2-51 — 








Copper, 50 p.p.m. 

4*64 — 








Copper, 10 p.p.m. 

4*32 6*38 








Zinc, 50 p.p.m. 

0*94 — 








JZinc, 10 p.p.m. 









Manganese, 50 p.p.m. 

0*98 — 







Manganese, lO p.p.m. 

2*77 — 







■ — 

Iron, 50 p.p.m. 

1-58 — 


1 15 






Iron, 10 p.p.m. 

2-29 — 








Cu X Zn 

2*84 — 








Cu X Mn 

371 -- 

, — 



— " 




Gu X Fe 

4*10 — 








Mn X Zn 

172 — 







— ; 

Znx Fe 

1-32 — 



— , 




, — 










Cu x Zn X Fe 

4-26 — 








Cu x Fe X Mn 

4'o6 — 









1*90 — 








Cux Mnx Zn 

3*59 — 




0 - 2 - 0 -I 




All metals (error) 

379 — 







The effect of copper, manganese and molybdenum 

The next experiment was concerned with evaluating the effect upon 
growth of different concentrations of copper and manganese and also of 
molybdenum, for reasons previously mentioned. A modified Belmont 
medium was devised in which the peptone was replaced by ‘Pronutrin’ 
casein hydrolysate (Herts Pharmaceuticals Ltd.) of equivalent nitrogen 
content and to which o-ooi % tryptophan was added. The resulting 
medium was much paler in colour than Belmont medium, so that the 
observation of pigment production was facilitated. Copper and manganese 
were added in the same form as previously used at the rate of o, 2, 4, 8, r6, 
32 and 64p.p.m. and molybdenum at 0, 2 and 4p.p.m. as sodium 
molybdate, ‘Analar’. These additions were made in all possible combina- 
tions. The media were dispensed in 10 ml. quantities in 50 ml. Pyrex 
conical flasks. Inoculation was from a suspension of washed spores grown 
on normal Belmont medium. 

Observations were made on the growth after 7 days’ incubation and on 
the material harvested after 10 days’ growth. The amount of surface growth 

Nutritioml requirements of Spikbury 215 

and the degree of darkening of the medium, as recorded in purely arbitrary 
units, is illustrated in Text-fig. 3. No significant difiference could be 
detected due to different molybdenum content, and the curves shown are 
therefore compounded for three observations for each point represented. 
It is obvious that both copper and manganese, but especially the former, 
had a marked effect on coverage, which was more than doubled by the 
addition of the optimum concentration of copper {c, 8 p.p.m.) to the 
control lacking the added metal. Increasing the addition of copper above 
this level resulted in a return to poorer coverage. The manganese effect 
was opposite to that of copper, increasing its concentration resulting in 
poorer coverage. Darkening of the medium was noticeably affected by 

Text-fig, 3. The effect of copper and manganese on surface growth and pigment 
production in a casein hydrolysate medium. 

both metals. In the absence of copper it retained its original colour, but 
the addition of increasing quantities of copper resulted in progressive 
darkening of the medium until with concentrations greater than 8 p.p.m. 
it became almost black. Again the manganese effect was directly opposite, 
its absence resulting in the maximum manifestation of the effect due to 
copper and increase of its concentration resulting in a progressively paler 
coloured fluid. Assuming that the darkening of the medium is correlated 
with some oxidase activity of the organism it seems likely that the copper 
and possibly the manganese play some role in the elaboration or physio- 
logical function of such an enzyme as has already been demonstrated in 
the mushroom by Keilin and Mann (1938). The mycelial dry weights are 
given in Table 2, and Text-fig. 4 shows the main effects of the metals 
plotted separately. Their statistical significance is apparent from Table 3. 


Transactions British Mycological Society 

Table 2. Effect of trace metals in a casein hydrolysate medium. 
lo days’ growth. Mycelial dry weights in mg. 

— indicates negligible growth. 

Added copper 

Added manganese concentration (p.p.m.) 

K ! 













Molybdenum absent 
31*1 27*0 34*2 
















































— ■ 

— _ 




Added molybdenum 2 p.p.m. 
26*8 28*9 23-8 






















































Added molybdenum 4 p.p.m. 
44‘i 22-2 25*4 










































35* X 








Table 3. Analysis of variance of mycelial dry weights of Streptomyces griseus 
grown on casein-hydrolysate medium with different trace-metal concentrations 

Main effects : 

Sum of 

Degrees of 









3* XI 












— ■■ 

First order interactions: 

Cux Mn 




", ■ 





■ ■ ' ; „ 


Mn X Mo 





' 0*2 

Second order interaction 





Nutritional requiremmts of SpUshury 217 

These data confirm the observations that copper, and, to a lesser extent, 
manganese are of significance in the development of the mycelial mat* 
Assays of streptomycin activity on selected crude filtrates from this experi- 
ment tended to confirm that the best titres were obtained with 8-16 p.p.m. 
copper and an equivalent amount of manganese. 

Parts per million of metal 

Text-fig. 4. The effect of copper, manganese and molybdenum on mycelial 
dry weights on casein hydrolysate medium. 

The effect of the nitrogen source on the action of trace metals 

Since it was apparent from the two preceding experiments that the 
growth of the organism was less vigorous upon 'Pronutrin’ than upon 
peptone, and that growth was modified in each case by the trace-metal 
content of the medium, it was thought desirable to devise an experiment 
to determine whether the nitrogen source modified the trace-metal effect. 
Belmont medium base was utilized, with the addition of 0*005% tryptophan 
and 0-00245 % cystine, the nitrogen being supplied from various sources 
at the rate of i mg. /ml. The growth of Streptomyces griseus on these media 
after 7 days’ growth is expressed in arbitrary units in Table 4. It will be 
noted that in some cases growth was very poor when copper and man- 
ganese were added, whilst in others good growth was obtained, and it is 
likely that, since no steps were taken to remove trace metals from the media, 
they were actually present in some of the nitrogen sources in sufficient 

21 8 Transactiom British Mycological Society 

quantity to permit growth to take place. Photographs of part of the 
experiment are shown in Pis. XII and XIII, and it will be noticed that the 
appearance of the mycelium is markedly influenced by both the nature of 
the nitrogen source and the concentration of trace metals added. 

Two facts are evident from this experiment : that the tolerance of the 
organism for copper and manganese is largely dependent upon the nature 
of the nitrogen source, and that there is in every case an optimum concen- 
tration of the trace metals for growth. This latter is more easily noted in 
Pis. XII and XIII, which were taken when the cultures were i o days old, than 

Table 4. Growth of Streptomyces griseus on different nitrogen sources 
with differing trace-metal concentrations. 7 days’ growth 

+ indicates amount of growth, t.= trace, st. = slight trace, — =no growth. 

Added Cu and Mn concentration (p.p.m.) 

Nitrogen source 






32 ‘ 



-f + + + 

■+• + + Hh 

+ + + + 

+ + + + 

4. 4- 4- 4- 



* Casydrol ’ 

+ + + + 

+ + -f + 

-j- -f + + 

+ 4 + + 

4 4 4 4 




+ + + 

+ + + 

+ + + 

4 + 4 





+ + + 

+ + + 

+ + + 





Ammonium nitrate 

+ + + 

+ + + 

+ + + 

-p 4. 4. 



Ammonium chloride 


+ + 

4- +■ 





Ammonium sulphate 

+ + 

+ + 

+ + 

4 4 





+ , 

+ + 












— - 









Potassium nitrate 








in Table 4, The increased tolerance of the organism towards trace metals in 
such media as those containing amino-acids and asparagin suggests thatyin 
these media, the copper and manganese is combined in organic form and 
is not active as a toxic cation. This toxicity is most readily observed in the 
presence of ammonium salts and other sources of nitrogen not readily 
available to the actinomycete. 

The major mineral constituents 

The effect of the so-called trace elements having been demonstrated it 
remains to discuss the effect of the major mineral nutrients. The empirical 
combination of mineral salts in Belmont medium had resulted in a medium 
superior in streptomycin and growth-producing properties to other media 
examined, and the following experiment was designed to assess the value 
of the different salts employed and to supply information relating to the 
nitrogen source and certain growth factor requirements, which will be 
discussed below. 

The experiment was designed factorially, using i % glucose as the carbo- 
hydrate source, and incorporated two nitrogen sources, ‘" Pronutrin’ casein 
hydrolysate and asparagin at a nitrogen level of i mLg.fmL; o-ooi % 
tryptophan and o*oor % cystine, and a control lacking these growth 
factors; 10 p.p.m. copper and manganese as ' Analar’ sulphates and a 

Nutritional requirements of Streptomyces griseiis. Spilshury 219 

control; three growth-promoting natural extracts and a control, and the 
following combination of mineral salts: 

{a) Control, no salts added. 

{b) NaNOg, 0-4%. 

{c) KH2PO4, 0*1 %. 

(i) MgS04, 0*05 %• 

{e) NaNOg -f KH2PO4, concentrations as above. 

(/) NaN03 + MgS04, concentrations as above. 

{g) KH2P04 + MgS04, concentrations as above. 

(A) NaN03 4- KH2PO4 + MgS04, concentrations as above. 

Because of the large size of the experiment the usual receptacles were 
abandoned and the media were distributed in 10 ml. quantities in 25 ml. 
glass bottles with narrow necks. After inoculation in the usual way the 
bottles were sloped so that as large an area as possible of the medium was 
exposed on which growth could take place. 

Growth observations were made at intervals, and after 10 days’ growth 
the material was harvested and the mycelial weights obtained. The 
beneficial effect of copper and manganese on the appearance of the growth 
was again confirmed, the presence of these metals giving a better surface 
growth than the control. Thick white growth was especially noticeable on 
the casein hydrolysate medium in the presence of phosphate and mag- 
nesium. Absence of the trace metals resulted in a pronounced slimy and 
submerged growth on ‘Pronutrin’ and, to a much lesser extent, on 
asparagin. On the latter medium the effect of the trace metals was to 
produce a grey and scaly growth rather than a white and powdery one. 
Very little growth took place in the absence of mineral salts. The presence 
of sodium nitrate alone resulted in a slight toxicity which was removed by 
phosphate and magnesium sulphate, particularly in combination: These 
latter were also stimulatory to surface growth together, but not singly. The 
appearance of all these combinations was influenced by the nature of the 
extract added and resulted in a great variety of growth forms. Some of 
these forms had already been observed on batches both of Belmont medium 
and some other media developed by Ainsworth et al, (1947). 

The mycelial dry weights are presented in Table 5, with their analysis 
of variance in Table 6, and it will be seen that the effect of the mineral 
constituents on dry weight is similar to that on growth as judged by 
coverage and appearance, and is highly significant when compared with 
other factors. This experiment emphasizes the need for a well-balanced 
medium with respect to all the factors, for the optimum development of 
S, griseus. 

The nitrogen source 

The organism appeared to be able to utilize a wide range of nitrogenous 
material, and many experiments were conducted in these laboratories in 
an endeavour to discover a suitable substitute to peptone in Belmont 
medium with varying degrees of success. It has already been shown 
(Table 4) that the best sources of nitrogen for growth appear to be 

220 Transactions British Mycological Society 

substances possessing amino groups, and that nitrates, ammonium salts, 
and nitrogen in cyclic molecules are less readily available. In addition to 
the substances listed above, aspartic and glutamic acids and glutamine 
provide a substrate suitable for normal growth. It must be emphasized 
that the availability of any particular substrate depends upon the presence 
and adequate concentration of the necessary mineral nutrients and growth 

In the experiment detailed in Tables 5 and 6 it is shown that i mg. /ml. 
nitrogen as casein hydrolysate gives a greater weight of mycelium than an 

Table 5. Weights of freeze-dried mycelium from different media 
{expressed in mg.) 

Added Cu and Mn 

Without added Gu and Mn (lo p.p.m. each) 











+ Tryp- 


+ Tryp- 

4 - Tryp- 









Control cystine 

Control cystine 

Control cystine Control cystine 

No salts, no extract 









I % meat 









I % yeast 









I % dung 









NaNOs, no extract 




1 6*8 





I % meat 









I % yeast 







1 8*2 


I % dung 









KH2PO4 , no extract 









I % meat 









I % yeast 


















MgS04 , no extract 









I % meat 








t 47 

I % yeast 





I 3 *t 




I % dung 









NaN03 4KH2P04, no extract 

1 7-1 








I % meat 

1 8*4 








I % yeast 









I % dung 









NaNO^-f-MgSO*, no extract 









I % meat 






49 *f> 



I % yeast 







■■TI:*4 . 


I % dung 



1 8*0 






KH2P04-f-MgS04, no extract 









I % meat 









I % yeast 



I 7‘3 



i 8 *o 



I % dung 









All salts, no extract 









I % meat 









I % yeast ^ ^ ^ 









I % dung 









Nutritional requirements of Streptomyces griseiis, SpUsbury 221 

Table 6. Analysis of variance on data in Table ^ 


Sum of 

Degrees of 






A (Nitrogen source) 

Main effects 
965*65 I 




X (Trace 'metals) 




— - 

a (Tryptophan and cystine) 




a (Mineral constituents) 






I (Growth factors) 







First-order interactions 
91*61 I 







> 0*2 






— , ,■ 







Xa , 

































5 1** 1*5 

• — 



Second-order interactions 
368*46 21 





1 8*44 


— ■ 

Xac & 













































— ■ 


Third-order interactions 
3094*92 7 







. — 












^ — ■ 









3118*15 21 



equivalent concentration of nitrogen as asparagin; yet there is no question 
that both are utilized by the actinomycete, and the evidence from this and 
from data presented below suggests that the most readily available nitrogen 
is that in amino groups. 

The effect of utilization of pure amino-acids as the nitrogen source 
as compared with ammonium nitrate is illustrated in Table 7. Belmont 
medium was used in which the peptone was replaced by i mg. /ml. 
nitrogen as indicated in Table 7 and to which o*oo8 % tryptophan was 
added. The experiment was carried out in quadruplicate with regard to 
nitrogen source, there being a control and sections incorporating o-ooi % 
concentrations of creatine, guanidine hydrochloride, and inositol which 
were added as possible growth factors. Actually they affected neither 

222 Transactions British Mycological Society 

growth nor titre, and the assay figures and the growth notes are therefore 
the means of four observations. For growth, straight-chain acids were 
better than those with branched chains, the very noticeable exception 
being histidine. The amide of the dicarboxylic glutamic acid is better than 
the monocarboxylic acids, and the only one giving growth comparable 
with ammonium nitrate. Branched chain acids gave weaker and browner 
growth and those containing the benzene ring very poor growth. _ Methio- 
nine was too toxic to permit growth, and was comparable in this respect 
with cystine. For streptomycin production the straight-chain acids were 
the best, but the branched chain ones gave higher titres than would be 
expected from the feeble growth produced. Histidine and glycine give 

Table 7 . The effect of different amino-acids on growth and 
streptomycin titre 

Nitrogen source 

Growth at 7 days 

Titre at 7 days 


Ammonium nitrate 

Good coverage 



Mycelium white with a mealy appearance 



Medium to good coverage, but mycelium 
tends to become brownish 

» >> 





j> jj 



Coverage decidedly poor, mycelium scanty 



and brownish 






» i) 





Very poor growth, mostly submerged 



)> >? 



Toxic; growth nil 


unexpectedly low titres, and also the titre for glutamine was less than that 
for alanine. Acids containing benzene rings, as would be expected, gave ! 
the lowest titres. , 

Several points of importance emerge from the above data. Organic 
nitrogen is not necessary for streptomycin production; nevertheless, with 
some organic nitrogen sources streptomycin can be produced even when 
the quantity of mycehum present is small. It has been shown that, in 9 
general, assuming trace metals to be in adequate concentration the titre 
follows closely upon the mycelial weight, and the conclusion can be reached 
that the Belmont medium owes its superiority over other media tested to 
its balance between the amino-acids of the peptone or papain digest and 
its mineral constituents. It is also obvious that different amino-acids fulfil 
different functions within the organism; for example, glutamic acid forms 
a good source of nitrogen for mycelium building, alanine favours strepto- 
mycin production although not promoting good surface growth, and i 

tryptophan, as will later be shown, is important in the initiation of growth. ^ 


Nutritional requirements £?/ Streptomyces griseus; Spihbury 223 

Growth-factor requirements 

It was early realized from experimentation on modifications of the 
Belmont medium that the omission of meat extract resulted in considerably 
poorer growth by the organism with corresponding loss of titre, and also 
that excessive autoclaving of this medium was deleterious to growth and 
could even result in complete failure of the inoculum to germinate. Several 
experiments were conducted with a view to ascertaining whether any 
accessory factors were necessary to the organism for its healthy growth in 
addition to the nutrients already mentioned. 

Tryptophan, known already to be essential to the development of 
Corynebacterium diphtheriae from the work of Mueller (1935) and others, was 
suggested as a possible requirement for Streptomyces griseus by poor growth 
on acid-hydrolysed nitrogenous material and the dying out of the organism 
on repeated subculturing on the asparagin medium of LePage and 
Campbell (1946). An experiment was arranged in which the relatively 
impure casein hydrolysate, "^Pronutrin^ and pure ‘vitamin-free’ casein 
hydrolysate (supplied by Ashe laboratories) were used as the nitrogen 
sources and to which tryptophan and cystine were added at the rate of 
50 and o-i mg./mL, respectively. The scope of the experiment was further 
extended by the addition of 5 % extracts of meat and yeast and an extract 
obtained by heating 500 g. of fresh horse dung with i 1. of water with 
constant mechanical stirring for J hr. at 80° C., filtering through glass-wool 
to remove the large particles and candling. This extract was dark brown in 
colour and will subsequently be referred to as ‘dung extract’. These various 
extracts were added at nine concentrations commencing at 10 % v/v of 
medium and halving the concentration at each level. This medium was 
distributed in 10 ml. portions in 50 ml. ‘Pyrex’ conical flasks, all the 
precautions customary when dealing with substances of a vitamin-like 
nature being observed in both this and all experiments subsequently 
described under this section. Inoculation was with o*i ml. of a washed 
spore suspension in 1/10,000 ‘Calsolene’ from a slope culture on LePage 
and Campbell’s medium. Growth after 10 days’ incubation is recorded in 
arbitrary units in Table 8. It will be readily observed that on ‘vitamin- 
free’ casein hydrolysate medium no growth took place in the absence of 
tryptophan and cystine, and that there was no factor in either of the 
extracts employed capable of replacing these substances. In the presence 
of both tryptophan and cystine, good growth was obtained, the amount 
of growth depending on the added extract as seen in Table 8. With 
‘ Pronutrin’ growth was obtained without added tryptophan and cystine, 
although it was of a submerged character and a true mycelial mat was not 
formed as on the previous medium with these additions. On adding these 
substances to the ‘Pronutrin’ medium growth was greatly enhanced. The 
mycelial dry weight of some of the treatments is^ shown in the histogram 
in Text-fig, 5. It may be concluded that tryptophan and cystine or one of 
these substances is important for the growth of S. griseus^ dcnd that some 
factor or factors present in extracts of meat, yeast and dung result in 


Transactions British Mycological Society 

Table 8. The effect of tryptophan and cystine and various natural extracts on the 
growth ^Streptomyces griseus on casein-hydrolysate media. 7 days’ growth 

t. = trace, St. = slight trace, no growth. 

Vitamin free casein hydrolysate 

Vitamin free casein hydrolysate 
+ tryptophan and cystine 


Prontitrin+ tryptophan and 

% extract 
added (v/v) 

Meat extract 

Yeast extract 

Manure extract 





~ , 









. _ 






















+ + 

4 4 

4 4 




4 4 4 4 



4 4 4 4 



+ + 




+ + -f + 

4 4 4 



4* + "f 




44 + 


4 4 


4 4 4 


4 4 







44 4 4 



4 4 

4 4 



















4 4 4 44 






4 4 








4 - 4 

4 4,^ 

■ 4 . 



4 4 

4 4 




4 4 4 


4 4 4 4 


4 44 


4 4 4 4 

4 4 4 



4 4 4 

4 4 4 

_l [- 




4 4 


4 4 

+ 4 

4 4 




4 4 



4 4 

4 4 



4 4 

. "h' 4: 

♦ All growth in this section was of the submerged type. 

improved growth. The small amount of tryptophan and cystine possibly 
present in ‘Pronutrin’ appears to be sufficient to permit growth but not of 
a normal character, and it is interesting to speculate whether this effect is 
in any way comparable with the submerged growth resulting from the 
unbalanced mineral nutrition which has previously been discussed. 

The need for tryptophan has also been demonstrated on a medium 
containing ammonium nitrate as the sole nitrogen source. It has also been 
found possible to replace tryptophan by a mixture of alanine and indole-3- 
butyric acid, and slight growth took place using alanine alone, although 

Nutritional requirements of Streptomyces griseiis. Spthhury : ^ ,225 

this may have been due to a small amount of tryptophan present in the 
alanine as an impurity. The effect of these substances are illustrated in 
PL XIIL This is of interest as suggesting that S* griseus is able to synthesize 
the indole ring but is apparently unable to synthesize aminoproprionic 
acid when provided with an inorganic nitrogen source, but it fails to explain 
the complete absence of growth on vitamin-free casein hydrolysate medium 
and further work is required to elucidate this point. 

An experiment was also carried out to demonstrate the effect of trypto- 
phan, cystine and dung extract upon growth when the nitrogen was 

Text-fig. 5. The effect of various growth factors on mycelial dry weight on 
caseift hydrolysate medium: lo days. 

supplied as asparagin. It should be explained that the asparagin used was 
from a natural Source and probably contained a small quantity of trypto- 
phan. The asparagin was incorporated in a basic Belmont medium at 
I mg. /ml. total nitrogen, and the following levels of substances were added 


• .. ,A - 

Tryptophan (w/v) 0*008 0*004 0*002 o*ooi 0*0005 o*o 

Cystine (w/v) 0*02 o*oi 0*005 0*0025 0*00125 0*0 

Dung extract (v/v) 1*00 0*5 0*05 0*0 — • — 

The main effects of these treatments are plotted as arbitrary units based 
on 2 days’ growth in Text-fig. 6, and it is seen that coverage is greatly 
increased by o*ooi % tryptophan and 0*5% dung extract. Increasing 
quantities of cystine inhibited good surface growth. As growth proceeded 
the differences between the treatments became less apparent. It was found 
that there was no significant difference in weight at different tryptophan 
levels after 10 days’ growth, and it would seem that tryptophan is only 
required in the earlier stages of growth. This was borne out by general 
observations made on other experiments. For example, when large inocula 
were used, sufficient tryptophan seemed to be carried over into the experi- 



226 Transactions British Mycologiccd Society 

mental medium from the inoculum either in the inoculum medium or the 
mycelium itself to permit inauguration of effective growth, a phenomenon 
similar to that encountered in yeasf-by Wildiers (1901), whereas when 
small inocula were employed and care was taken to use spores which had 
been carefully washed, growth did not proceed on media lacking trypto- 
phan. Thus in the experiment quoted above relating to the synthesis of 
tryptophan it appears likely that there was sufficient tryptophan present 
to initiate slight growth, since a very small growth took place in the control 

Text-fig. 6. The effect of tryptophan, cystine and dung extract on surface growth: 2 days. 

Asparagin us^ as the nitrogen source. 

lacking added tryptophan and alanine. The depressing action of cystine 
and the stimulatory effect of dung extract were, however, reflected in the 
mycelial dry weights. 

To sum up, it has been shown that tryptophan is required during the 
early period of growth from the spore, that cystine is not essential and 
proves inhibitory in quite small concentrations, and that there are certain 
unidentified factors present in extracts of meat, yeast and horse dung 
which are valuable auxiliaries to a healthy growth. According to Schopfer 
(1943) little seems to be known of the growth factors required by actino- 
mycetes. It was suspected that creatine, guanidine, or inositol might act 
as growth factors, but tests on these substances proved negative. Other 
substances investigated in low concentration by Miss E. Brookfield, of a 
type known to influence growth in higher plants were indole-3-acetic acid 
indole-3-butyric acid, and naphthaleneacetic, naphthoxycetic, phenoxy- 
acetic, 2, 4, dichlorophenoxyacetic and 2, 4, dichloropropionic acids. These 
had either no effect on growth or a depressing one. 

Nutritional requirements of Streptomyces griseiis* Spikbury 227 


Evidence is presented relating to the essential constituents of a medium for 
optimum growth ot Streptomyces griseus in surface culture and the production 
of streptomycin by this organism. 

A comparison between various inorganic and organic sources of nitrogen 
is made with glucose as the carbohydrate source, and observations relating 
the growth on these substrata with the mineral nutrition of the organism 
are included. Various trace elements, are demonstrated to be important, 
notably copper, manganese and zinc; and the addition of sodium nitrate, 
potassium phosphate and magnesium sulphate are shown to be beneficial. 

Tryptophan is shown to be essential in the early stages of development, 
and the stimulatory effects of various other adjuvants are discussed. 

I should like to express my thanks to Dr B. C. J. G. Knight of these 
laboratories for suggestions relating to the growth factor requirements of 
Streptomyces griseus and for independent confirmation of the effect of 
tryptophan on the early growth of the organism; to Dr G. C. Ainsworth 
for his continual interest and helpful suggestions throughout the progress of 
the work; and to Miss E, Brookfield for the communication of some of her 
experimental results recorded in this paper. 

My gratitude is also extended to all the members of the staff of these 
laboratories who took part in the experimental work, especially my 
assistant, Miss H. Bishop; Miss E. Howard who was responsible for the 
assays, and Miss G. Puddefoot who performed most of the work of 
statistical analysis. 


Ainsworth, G. C., Brown, Annie M., Marsden, P. S. F., Smith, P. A. Sc Spilsbury, J. F. 
(1947). A method for the production of streptomycin in surface culture. J. Gen. 
Microbiol, i, 335-43. 

Brown, Annie M. Sc Young, P. A. (1947) . A dilution method for the assay of strepto- 
mycin. J. Gen. Microbiol, i, 353-60. 

ELeilin, D. & Mann, T. (1938). Polyphenol oxidase, purification, nature and properties. 
Proc. roy. Soc. B, cxxv, 187-204. 

LePage, G. a. & Campbell, E. (1946). Preparation of streptomycin. J. bioL Chem. clxii, 

Mueller, J. H. (1935). Studies on cultural requirements of bacteria. J. Bact. 

Richards, F. J. (1941). The diagrammatic representation of the results of physiological 
and other experiments designed factoriaily. Ann. BoL, Lond.y N.S., v, 249-296. 
Robbins, W. J. Sc Hervey, A. ( 1 944) . Response of Pythiomorpha gonapodyides to manganese. 
Bull. Torrey bot. Cl. lxxi, 258-266. 

Schopfer, W. H. (1943). Waltham, Mass., U.S.A. 

Stiles, W. (1946). Trace Elements in Plants and Animals. London. 

Wildiers, E. (1901). Nouvelle substance indispensable au developpement de la levure. 
Ce/Me, xvin, 31^31. 


228 Transactions British Mycological Society 

Plate XII 

Fig, I A. Strepiomyces griseus, lo-day growth. Reading from top to bottom and 

: left to 

Nitrogen source 
"Ammonium nitrate 


p.p.m. trace metals 



Ammonium chloride 




A - 

Ammonium sulphate 













Plate XIII 
Fig. IB 





1 Guanidine 




B < 











.Evans’s peptone 




Fig. 2, Strepiomyces griseus. 7-day growth. Reading from left to right. 

Control, Ammonium nitrate without growth factors. 

Ammonium nitrate + 0-005 % dl-tryptophan. 

Ammonium nitrate +0-002 % alanine +0*005 % indole-3-butyric acid. 

Trans. Brit. Myc. Soc, 

[ 229 1 


By C. H. GADD and C. A. LOOS 

Tea Research Institute of Ceylon 

(With I Text-figure) 

Exobasidium vexans was first described by Massee (1898) from material sent 
by Dr Watt from Assam, where it was causing a serious disease of tea termed 
^Blister Blight’. The disease was well described by Watt and Mann (1903) 
and was later studied by McRae (1910) and Tunstall and Bose (1921) in 
North India, Du Pasquier (1933) in Indo-China, and by Sawada (1922) 
in Formosa and Japan. 

The disease becomes evident on young expanding leaves as translucent 
spots which later become conspicuous, circular ^ blisters The upper surface 
of the leaf becomes indented to form a circular pit, and the lower surface 
protrudes to form the so-called blister. A few days later the convex surface 
of the blister becomes white, with a velvety surface, as the fungus is revealed 
by the splitting and removal of the cuticle and epidermis. 

This disease appeared for the first time in Ceylon in October 1946 and 
rapidly spread through a large area of tea over 1500 ft. above sea-leVel. 
The symptoms leave no doubt that the disease is the same as that described 
from North India, and in our opinion the causative fungus is undoubtedly 
E. vexans Massee. 

In his description of £. vexans^ Massee described two kinds of spores: 
(i) small non-septate basidiospores, and (2) large one-septate conidia. 
Sawada (1922) however put a very different interpretation on his observa- 
tions, He wrote: 'The present writer believes, however, that the so-called 
conidia borne on the convex under-surface of the blistered area are really 
bicellular basidiospores. He has failed to find any spores on the tips of 
simple conidiophores as described by Massee and McRae. On following 
out the development of the basidiospores he found that the unicellular 
spores borne on the basidium may fall from the sterigmata when ripe or 
may remain attached until the basidia wither and collapse; in either case 
they ultimately develop a septum,’ 

The occurrence of the disease in Ceylon gave us an opportunity* to study 
this fungus with special reference to the production , of basidiospores and 
conidia. Our observations lend support to Sawada’s view and it is thought 
that they will be of interest to other students of this group of fungi. 

The fungus. The velvety appearance of the convex surface of the blister, 
when it first becomes white, is due to the presence of dense clusters of 
vertical sterile hyphae, approximately equal in length, each with a rounded 
apex. Some of the basidia, if not all, project above this layer and can 
easily be recognized by the sterigmata (Fig. i). 

In his description, Massee states ' Some of these hyphae run out into 

230 Transactions British My cological Society 

long sterile filaments giving a minutely downy or velvety appearance to the 
blister when seen under a lens; the great majority of the hyphae, however, 
remain short and produce a single conidium at the apex’. In his figure of 
the hymenium, he shows three basidia projecting above a cluster of 
conidiophores, each conidiophore bearing a thick-walled two-celled 
spore, but he does not figure any sterile hyphae which give the surface its 
velvety appearance. Watt and Mann, and Du Pasquier reproduce Massee’s 

McRae (1910) figures a cluster of vertical hyphae above which one 
basidium projects slightly. At the apex of some of the vertical hyphae are 
figured two-celled thin-walled conidia. The conidiophores are of about the 
same length as the sterile hyphae, not shorter as described by Massee. 

Fig. I. A, Thin-walled basidiospores, B, Thick-walled basidiospores. G, Basidium with three 
sterigmata. D-G, Basidia and their basidiospores. H, Basidium with a septum near 
apex. I-N, Germinating basidiospores. O, Transverse section through a young blister 

Tunstall and Bose (1921) give a dia^ammatic representation of a section 
through a diseased leaf showing vertical conidiophores and sterile hyphae, 
the former being distinguished by the presence of solitary one- or two- 
celled thin- walled spores at their apices. In this respect their interpretation 
agrees with McRae’s. 

Conidia, Massee described the conidia as "hyaline or with a tinge of 
yellow when seen in the mass, elliptic with somewhat pointed ends, ■ 
one-septate, slightly constricted at the septum, straight, or sometimes very 
slightly curved, measuring 14-16 x 5-6 /x’. Tunstall and Bose describe 
them as usually single cells, but occasionally uniseptate, measuring 
1 1-2 1 X 4*5-6-5/^. Du Pasquier wrote "Les conidies sont les plus abondantes. 
L’extremite renflee du filament grossit, se separe de la tige par une cloison, 
et forme une spore amincie aux deux extremites unicellulaires ou parfois 

Exobasidium vexans. C. H. Gadd and C. A. Loos 231 

^ uniseptte, qui mesure 1 1-20 x 4- 5-6-5^.’ Mann (1906) and McRae 

describe the spores as two-celled but give no measurements. 

There is thus fair agreement regarding the size of the spores, but Tunstall 
and Bose and Du Pasquier agree that they are usually non-septate. We 
have found very few non-septate spores except those attached to basidia. 
Two-celled spores are abundant, free on the surface of the blister, and they 
are undoubtedly the spores commonly referred to as conidia. 

Basidiospores. The basidia are loosely distributed among the sterile 
hyphae (or conidiophores) and often project somewhat above their level. 
I They are sub-cylindric, usually with two slender spine-like sterigmata 

(Fig. I D-G), though occasionally three have been seen (Fig. i G). 
A septum sometimes occurs near the apex (Fig. i H). 

Massee described the basidiospores as ‘hyaline, continuous, glabrous, 
ovate, oblong, often slightly inequilateral, 5 x 3ja’. Mann writes of them 
as ‘extremely minute oval bodies borne in pairs at the ends of special 
erections termed basidia\ Tunstall and Bose give much larger measure- 
ments, 7~'i3*5 X 2*3~4*5/x and describe them as ‘sausage-shaped’. Du Pas- 
quier gives Tunstall and Bose’s measurements with Massee’s figures, and 
the following description ‘Les basidiospores n’apparaissent qu’a certains 
moments et le plus souvent au debut de la saison. L’extremite du filament 
se ramifie en 2 a 5 comes effilees qui portent les spores. Celles-ci sont plus 
petites que les precedents (7-13 x 2*3-4'5/x), hyalines, cylindriques, 
arrondies aux deux bouts.’ 

The small spores described by Massee and the larger non-septate basidio- 
spores of Tunstall and Bose are almost invariably attached to sterigmata 
^ and leave no doubt that the smaller ones are an earlier stage of the larger 

ones. Our measurements of the larger non-septate spores gave a range Oa 
6-14x2*8-4*3//. which, with their shape, indicates that they are the 
basidiospores of Tunstall and Bose. 

At this stage our observations indicated that the basidiospores were 
non-septate, but no explanation could be ofiered of the origin of the more 
numerous septate spores lying on the surface of the blister. There was no 
evidence that the basidiospore enlarged and became septate after detach- 
ment from the sterigmata, as suggested by Sawada. There was the possi- 
bility, however, that the spores might enlarge and become septate before 
detachment, but that hypothesis also carried the proviso that the septate 
spores must fall at the slightest disturbance. 

We were fortunate in having diseased bushes in close proximity to the 
' laboratory and were able to examine freshly gathered material, subjected 

to a minimum disturbance. In that material we found one-septate spores 
in situ on the basidia (Fig. i F, G) and although even small disturbance 
displaced them, sufficient were seen to afford clear proof of their origih. 
The two-celled spores attached to sterigmata measured 14-17 x 3*5-5*6/x. 
Measurements of similar spores lying on the surface of the blister (Fig. i A) , 
and previously regarded as the conidia of Massee, gave a range of 
13-27 x 4*3-6*5/x and a mean (of 50 spores) of 18*7 x 5*5jLt. 

It thus became evident that the larger basidiospores described by Tunstall 
and Bose are not fully grown, and that the spores lying free on the surface 

232 Transactions British Mycological Society 

are in fact mature basidiospores and not conidia as was previously supposed. 
The unicellular conidia reported by Tunstall and Bose were probably 
displaced basidiospores though it is somewhat surprising that they were 
more abundant than the septate form. 

Interputation. The ease with which septate basidiospores are dislodged 
I affords a simple explanation of their abundance on the hymenial surface 

and their sparsity at the points of production, even in carefully handled 
materiaL Yet Massee and others in their figures have shown these spores 
being produced on conidiophores. Are we to assume that these authors, 
unable to connect the larger two-celled spores lying on the surface with the 
non-septate spores visible on sterigmata, concluded that they had some 
other origin and made an obvious but unjustified reconstruction? Sawada 
appears to think so, and we incline to that view, not only because of the 
negative evidence that they had not seen the two-cdled spores attached to 
conidiophores, but also because of the positive evidence connecting the 
non-septate basidiospores with the one-septate ‘conidium\ 

One further point needs to be mentioned. Massee's figures show his 
‘conidia’ to be thick- walled. Neither he nor later writers make direct 
reference to the ‘conidia’ being thick- walled. The two-celled basidiospores, 
so abundant on young blisters, are thin- walled, but in older material we 
have seen numerous free spores with thick walls (Fig. i B) similar to those 
illustrated by Massee. In the same material basidia and hyphae with 
thickened walls were also seen. It would seem that ageing, or a change in 
climatic conditions, may induce wall thickening. The size and shape of the 
thick-walled spores, coupled with the fact that none could be found on 
a conidiophore, left no doubt in our minds of their identity as basidiospores. 

Over-wintering. Th.t spores oi Exobasidium vexans are reputed to be short- 
lived, and no very satisfactory explanation has been offered of the manner 
by which this fungus survives unfavourable seasons and years. The 
thick-walled basidiospores remind one of resting spores, but whether the 
thickening of the wall increases the viability of the spore and enables it to 
survive during unfavourable conditions has not been determined. It is 
perhaps a hypothesis worth consideration. 

Germination. The two-celled basidiospores germinate readily in hanging 
drops of water, germ tubes emerging in about five hours (Fig, i I-N). 
Usually germ tubes emerge from both ends of the spore (Fig. i j), but 
occasionally an additional germ tube from a side wall has been seen 
(Fig. I L). As germination proceeds the spore begins to clear (Fig, i I, K), 
as though the protoplasm is streaming very slowly into the germ tubes 
leaving the centre of the spore empty. If only one germ tube is formed, 
only the half which forms the tube clears (Fig. i K). More commonly, 
both ends produce germ tubes and a circular clear area first becomes 
evident around the septum (Fig. i I). This area enlarges as growth 
proceeds until the spore is clear and apparently empty (Fig. i N). 

In such empty spores, not one but two and even three septa are some- 
times clearly visible, although before germination only the middle one was 
apparent. It will be evident that the additional septa are formed, as a rule, 
during germination, otherwise the flow of protoplasm from the centre into 

Exobasidiiim vexans. C. H. Gadd and C. A.. Loos 233 

the germ tubes would be prevented. Support is lent to this view by the 
fact that spores which produce germ tubes from side walls invariably have 
an additional septum, which is probably there when germination starts, 
as the only way by which the protoplasm can move from a middle com- 
partment is by pushing out a germ tube from the side wall. The fact that 
germinating spores can rapidly make cell-wall material and so form 
additional septa within the spores possibly supports the view expressed 
above that under certain conditions the spore is able to thicken its walls by 
the addition of more material. 

No haustoria have been observed either in hanging drops or on leaf 
surfaces. It is probable therefore, as stated by other authors, that entry into 
the leaf takes place through the stomata, although such entry has not been 
:seen by us. 


Blister Blight of tea, a leaf disease caused by the fungus Exobasidium vexms 
Massee, occurred for the first time in Ceylon in October 1946 and spread 
rapidly through a large tea area. Massee and others have stated that 
E, produces two-celled conidia as well as non-septate basidiospores. 

Sawada has dissented and expressed the view that the two-celled spores 
are in fact mature basidiospores. 

Observations described here support Sawada’s opinion. The mature 
two-celled basidiospores are very easily dislodged from the sterigmata and 
are usually to be found on the surface of the blister. In carefully collected 
material we have seen the mature two-celled basidiospores attached to 
:sterigmata, and so have demonstrated the origin of spores lying on the 
surface: they are mature basidiospores and not conidia. Under certain 
conditions the cell wall thickens and the spores then closely resemble 
Massee’s figures of ^conidia’. Neither Massee nor later writers make any 
mention of the 'conidia’ having thick walls, which must be interpreted 
that normally they are thin-walled as seen by us. Massee’s drawings of 
''conidia’ are not in error, though his reconstruction of their origin is. 

Germination of the spores is also described. Although the basidiospore 
is normally one-septate, as many as three septa have been seen in ger- 
minated spores. The view is expressed that the extra septa are normally 
formed during germination. 


Du Pasquier, R. (1933). Principales maladies parasitaires du ihiur et du cafeier en extime- 
orient, pp. i 65->9. Hanoi. 

Mann, H. H, (1906). Blister Blight of tea. BulL Indian Tea no. 3. 

McRAri, W. (1910). The outbreak of Blister Blight on tea in the Daijeeling district in 
igoS-g* BulL Indian Tea Ass, no. 3. 

Massee, 0. (1898). Tea blights. JT^sie; 1 09--1 1. 

Sawada, K. (1922). Gan Exobasidium vexans '^2^^. bear conidia besides the basidiospores? 

Trans, not, hist. Soc, Formosa, ux, 7, (Abstract Rev, appL MjcoL i, 454.) 

Tunstall, a. G. & Bose, S. C. (1921). The fungus diseases of the tea leaf. Quart, J, 
Indian Tea Ass. pp. aog^t^. 

Watt, G. & Mann, H. H. (1903). The Pests and Blights of the Tea Plant, 2nded. pp. 387-91. 

{Accepted for publication % 1947) 

[ 234 ] 


By E. J. H. CORNER, M.A., F.L.S. 

(With 9 Text-figures) 

Asterodon is a genus of Basidiomycetes with one species, A. ferruginosus, 
which grows on rotten coniferous wood in northern and montane parts of 
Europe and North America. Its brown resupinate fruit-bodies develop 

Fig. 1. Skeletal hyphae, generative hyphae and stellate setae from the fruit-body 
of Asterodon ferruginosus, X ^00, ^ ^ ^ ^ ^ ^ ^ ^ ^ ^ 

Notes on Asterostroma and Asterostromella, E. J. H. Corner 235 

Hjdnum-like spines which point vertically downward. Mycelial patches on 
the upper sides of fallen logs are sterile and without spines. Microscopically 
the tissue of the fruit-body (and, also, that of the mycelium) is peculiar in 
developing stellate setae with thick brown walls. This feature occurs in 
Asterostroma as well, but it has CorticiumAik^^ fruit-bodies without spines. 
Patouillard (igoo), therefore, made the group Asterostromeae, as a series 
of the Hydnaceae, to cover the two genera. Bourdot and Galzin (1928) 
treat the group as a subtribe, Asterostromaceae, of the Porohydnaceae. 
Donk (1933) has pointed out, however, that the group should be placed in 
the Hymenochaetoideae which he regards as a subfamily of the Aphyllo- 
phoraceae. Actually the position of the group cannot be decided until the 
extent of the Hymenochaetoideae or, as I prefer to call them, the Xantho- 
chroic Series is known. These xanthochroic fungi are distinguishable not 
so much by the Hymenochaete setae, which are absent from many of them, 
as by the characters of their hyphae. The absence of clamp connexions, the 
lack of inflation of the cells of the fruit-body, and the ochraceous or brown 
colour of the hyphal walls, which darkens to ferruginous or date brown 
with alkali, distinguish these fungi. Their fruit-bodies may be Clavarioid 
(as Lachnocladium)^ Hydnoid, Stereoid, Corticioid or Polyporoid; they have 
a prevailing yellow-brown colour and fibrous, coriaceous or ligneous 
texture; and they are built up entirely by apical growth (thus the Agaric 
form with its typical button stage is absent). Setae, dichophyses, skeletal 
hyphae, gloeocystidia and so on distinguish various genera, or subgenera, 
but, except as products of this kind of hypha, they are not unique, for 
similar organs occur in other series of Basidiomycetes. 

In this paper I wish to explain how this little-known fungus, which may 
be amenable to laboratory culture and experiment, has two features which 
should greatly assist in investigating the form factors, or evocators, of the 
fruit-body: they are the plastic growth of the setae and the direction of the 
skeletal hyphae. 

Description of Asterodon ferruginosus 
A. ferruginosus Pat., Soc. mycoL Fr, x, 1894, 129. 

Synonym: Hydnum ferruginosum Fr. fide Lundell, Sver. Svamp. fasc. vn-vm, 1936, 23. 

Synonymia alia vide Rogers et Jackson, jPar/. I, 1943, 271. 

Fruit-body resupinate, widely effused, forming a floccoso-felted ferruginous-ochraceous 
or tawny subiculum of very variable thickness, 0*2-8 mm. thick; margin bright tawny 
ochraceous, floccose, often tumid; spines i— 1*5 mm. long, o*2-o*3 mm. wide at the base, 
developed centrifugally, crowded in the centre, subulate, simple, subpruinose fawn- 

Gn very rotten coniferous wood: Sweden, Finland, Siberia (on Abies sibiricae a.nd Pirns 
silvestrisy fide FisincGi U.S. A. {on Pinus axxd Abies),* 

Spores 5-6 X 4-4*5 fz, or 6-8 x 4-5 fz (Comer, ReL Farlow. 302), white, smooth, ellipsoid, 

Basidia 18-25 x 5-8 pc, 14-20x6-746 (Corner, RsL Farlow, 302): sterigmata 4, 

Cystidia 16-25 x 3-546, subventricose with a narrow cylindric appendage 1*546 wide, 

* According to Pilat (1935), the specimen recorded on Populus by Bourdot and Galzin 
is Hymenochaete cirtnamomeum. 

236 Transactions British My cological Society 

• thin-walled, colourless, vacuolate, only in the incipient hymenium at the tips of the spines 
and at the margin of the resupinate part, soon collapsing, 

Hymenial setae 25-45 x 5- 10 ft, fusiform-sub ventricose, acute, straight or curved at 
the base, with smooth thick brown walls, mostly simple on the spines, usually suh stellate with 
I -*4 short branches in the intervals between the spines, 

Extrahymenial setae only in the tissue of the spines, 60-170x6-7^, straight, longi- 
I tudinal, simple or, rarely, once bifurcate (never stellate), sometimes with a short lateral 

branch transformed into a hymenial seta, aseptate, acute, with smooth brown walls to 
f 2*5 ju thick, with a longer or shorter stalk tapered to the generative h^^ha. 

Stellate setae only in tissue of the resupinate part of the fruit-body, with 4-6 acute arms 
to 90 ft long X 5“7ft wide at the base, rigid with smooth brown walls 1-2 ft thick, rarely 
once bifurcate, often somewhat curved, aseptate, the central body scarcely enlarged, the 
stalk to 100 ft long and tapered to the generative hypha : largest in the basal tissue next the 
substratum, smaller towards the subhymenium, and with all gradations to the substellate hymenial 
setae between the spines. 

Hyphae dimitic:* generative hyphae 1-5-3 ft wide, septate without clamps, with thin 
and colourless, or slightly thickened and pale ochraceous, smooth walls: skeletal hyphae 

• 2 *5-3 *5 ft wide, unbranched, or rarely bifurcate once, aseptate, unlimited, with smooth 

■ brownish ochraceous walls, 0*5-1 ft thick, the lumen distinct. 

Notes on Asterostroma and AsterostromeUa. H. Comer 237 

Strwtme of the fruit-body. The general construction is shown in Fig. 2, and 

details of me nymemiim in Figs. 3 and 4. The main features are the 
direction of the hyphae and the size, shape and direction of the setae. In 
the resupinate flesh, the hyphae are loosely interwoven in all outward 
directions from the substratum, but they have a horizontal centrifugal 
tmdency m the basal layer next the wood. The skeletal hyphae end 
obliquely in the hymenium or subhymenium or against the substratum 

Fig, 3. A longitudinal section of the tip of a spine of Asterodm ferrugimstts, x 500. 

and have simple cylindric tips. The tissue is denser towards the sub- 
hymenium because of the more frequent branching of the generative 
hyphae and the greater number of small stellate setae. The subhymenium 
itself, is poorly defined, 5-10/^ thick, and the hymenium is a continuous, 
unthickened palisade i6-2op. deep. In the spines the hyphae are strictly 
longitudinal, entwined but not interwoven: the skeletal hyphae are often 
kinked or irregularly flexed unlike those of the resupinate flesh. The 

238 Transactiom British Mycological Society 

density of the tissue of the spines is caused by the abundance of compactly 
longitudinal skeletal hyphae and the numerous extrahymenial setae 
among them. The tips of the spines are sterile and are composed of the 

down-growing ends of the three tissue-elements. 

Development of the fruit-body. Although I have not seen young specimens, 
it is clear that the resupinate part must develop as a superficial centrifugal 
outgrowth from the mycelium in the wood. It varies greatly in thickness 

Fig. 4. A section of the hymenium between two spine bases of Asterodon fsrruginosus, 

X 500; spores, x 1000. 

and is composed of hyphae which have at first grown outward in all 
directions, so as to be loosely interwoven, and are compacted by the arms 
of the stellate setae. At a certain stage of development the uniform out- 
growth slows down : the stellate setae become smaller, with shorter branches 
and more evident axis at right angles to the future hymenium, and the 
generative hyphae branch more freely. The downgrowth then stops: the 
end-cells of the generative hyphae become hymenial setae or basidia and, 
by sympodial intercalation of new basidia, the single-layered hymenium is 
constructed. In certain parts, however, or, indeed, over the whole central 
region of the incipient hymenium, there develops a Afield’ corresponding 
to the pore-field of polypores (Corner, 1932 in which the spines arise as 
localized areas of downgrowth o*2~o*3 mm. diameter. All the hyphal 
tips in such an area become positively geotrophic and, in the renewed 
excrescence, aline themselves to build the spine. Thus the formation of the 
spine is unidirectional in contrast with the multidirectional development of 
the resupinate flesh. Subsequently the hymenial factor develops on the 

Notes on Asterostroma and Asterostromella. E. J. H. Comer 239 

spine and makes the hyphal ends on the sides of the down-growing fascicle 
of hyphae turn out at right angles, as if diageotropic, to construct the 
hymenium. The spines are the positives of the tubes of polypores. 

The direction of the hyphae. As there is no inflation of the skeletal hyphae, 
which soon become more or less rigid with thickened walls, they suffer no 
displacement and their direction in the mature tissue shows the original 
direction of their apices, which in turn implies the direction of the force 
controlling their growth. In the basal layer of the resupinate flesh there 

appears to be a centrifugal force causing alinement of the hyphae against 
the substratum : it may be thigmotropic or diageotropic. In the rest of the 
flesh there appears to be no controlling factor other than an inherent 
divergence, as a general excrescence, which dies away until resuscitated by 
positive geotropism in the spine areas or terminated by the hymenial factor 
in the network surrounding the areas. Eventually a similar diversion of the 
hyphal tips on the sides of the spines, as occurs in the basal layer of the 
flesh, leads to the acropetal extension of the hymenium down the spines, 
the tips of which, however, are always sterile. This point and the fact that 
the hymenium never develops on upturned resupinate surfaces show, 
nevertheless, that the hymenial factor is in some manner geotropic as well 
as intrinsic. 

240 Transactions British Mycological Society 

The shape of the setae. It is, at first, puzzling that a fruit-body, which is so 
apparently simple, should have such a variety of hyphal endings. If it be 
assumed, however, that the setae are h^^hal ends of limited growth not 
entirely deprived of the faculty of branching, then it can be seen that their 
shapes express the form-factors of the fruit-body. In the loose tissue of the 
resupinate flesh, the stellate form with equal branches indicates the multi- 
directional outgrowth of the hyphae where there are no evident form- 
factors : that is, the stellate form is the uninhibited. As the outgrowth dies 
away, so the stellate setae become smaller. In the hymenium, where the 
hyphal ends no longer branch, the setae become simple and alined with 
the basidia. The parts of the setae which lie in the subhymenium become 
shortly branched to form the substellate setae of the hymenium between 
the spines. In the spines branching is suppressed and the setae are trans- 
formed into the elongate longitudinal extrahymenial setae, expressing the 
unidirectional parallel (not expanding) downgrowth, but where the 
peripheral extrahymenial setae come within the influence of the hymenial 
factor near the tip of the spine, they produce an out-turned branch which 
becomes a simple hymenial seta, as does any hyphal tip which becomes 
modified into a seta in this hymenium. In proof of this relation between 
the shape of the seta and the form-factors of the fruit-body, and as an 
indication of what takes place in the spine area on inception of a spine, 
one can analyse the shapes of the setae at the base of the spine, as shown in 
Fig. 5. Just before the spines develop, outgrowth in the spine area is 
stimulated and the tips of any hyphae in the area are aflfected; thus, the 
setae at the base of the spine have often reduced arms at the level of the 
hymenium between the spines and an elongate ray entering the base of 
the spine as one of the first-formed extrahymenial setae: such are, in fact, 
diminutive stellate setae of the resupinate flesh caught in the act of 
development and converted into setae of the tissue of the spine. The 
slightly oblique ray shown in Fig. 5, as entering the base of the spine on its 
right, has become caught in the hymenium and terminates as a hymenial 
seta as well as providing a branch. The corresponding seta on the left of 
the spine base, however, has just escaped the hymenium and been deflected 
into the tissue of the spine where it ends blindly. The two examples show 
that the influence of the hymenial factor, whatever it may be, is sharply 
delimited and that the narrow subhymenium, so difficult to examine and 
to define microscopically as often to be overlooked, expresses in its very 
nondescript appearance the neutral zone between the medullary tissue of 
the spine and its hymenial envelope. 

Relation between setae and skeletal hyphae. The two elements are always 
distinct in this fungus. Both arise from generative hyphae, as modified 
laterals, but always independently, the stalk of one never connecting 
directly with the stalk of the other. The setae have limited growth and 
become embedded in the tissue: the skeletal hyphae have unlimited growth 
and terminate at the surface of the fiuit-body. Both are thick-walled and 
aseptate, but the skeletal hyphae are only half as wide as the extrahymenial 
setae or as the arms of the stellate setae. Thus this fungus corroborates the 
impression gathered from the xanthochroic polypores (Corner, 1932^) that 

Notes on Asterostroma and Asterostxomella. E. J, H. Comer 241 

there are extrahyinenial setae, as well as skeletal hyphae, in the construe- 

tion of xanthochroic fruit-bodies. 

Experimental use 

If Asterodonferruginosus can be induced to fructify in artificial culture/ the 
shape of the setae and the direction of the skeletal hyphae under different 
experimental conditions should reveal how the hyphae have reacted. Thus, 
rotation on a klinostat, the effect of various radiations, or the action of 
gaseous or volatile chemical compounds could be studied in an attempt 
to find out what are the factors influencing hyphal growth and the 
development of hyphal tissues — matters which at present merely excite 
wonder. It should be possible, also, to find out by differential response 
what is the difference between setae and skeletal hyphae and, thus, what 
is the meaning of one of the distinguishing features of this series of Basidio- 

Asterostroma Mass. 

This genus has the stellate setae of Asterodon^ but its fruit-bodies are 
Corticioid without spines, the hyphae are monomitic (without skeletals), 
and there are no extrahymenial or hymenial setae (unless, perhaps, in 
A, degenerans Bres.) ; in their place most species have thin-walled colourless 
cystidia, containing large oil globules, which are homologous with the 
gloeocystidia of Lachnocladium, There are some twenty species, mostly 
lignicolous and, probably, in the main tropical. I have studied several 
species in Malaya and find their construction essentially as in Asterodon^ but 
simpler. The fruit-bodies have a thin basal layer of horizontal hyphae, 
spreading over the substratum, from which a loose cortex develops on the 
lower side as a vague multidirectional excrescence of hyphae with retarding 
growth : finally, this is covered by the even, single-layered hymenium. The 
largest stellate setae occur in the basal and initial region of the cortex and 
they decrease in size towards the hymenium, A. degenerans seems to have 
substellate setae transitional to typical Hymenochaete-sttdit^ thdiX is, with a 
few short basal branches. In a few species, as A, ochroleucum Bres., the 
branches of the stellate setae may be once dichotomous, or even 2-3 times 
(A. laxum Bres.) : in A. muscicolum (B, & C.) Mass, there appear to be 
irregular shapes suggesting transition to the dichophyses of Asterostromella. 
Some species, as that figured in Fig. 6, have small basidia; others have 
strongly projecting basidia as in Asterostromella, but always in a continuous 


This genus is composite and consists of species related with Aleurodiscus^ of 
true xanthochroic fungi, and, probably, of other Thelephoraceous deriva- 
tives. Its character is the presence of dichophyses, as thick-walled dichoto- 
mizing hyphae of limited growth, forming the mass of the tissue in the 
resupinate Corticioid fruit-body. I wish to indicate merely the xanthochroic 
species, several of which I have studied in Malaya. They resemble 
stroma in most points, particularly the monomitic construction without 



Notes on Asterostroma and Asterostromella. E. J. H. Comer 243 

clamp connexions, as shown in Figs. 8 and 9, The dichophyses are lateral 
branches of the generative hyphae, which are usually extremely tenuous 
and difficult to see: the dichophyses have limited growth, though the 
thinness of the fruit-body may obscure this fact, and in place of the stiff 
radiating arms of the stellate setae they become progressively dicho tomous 

Fig. 8. A section of the resupinate fruit-body of a xanthpchroic species of Asterostromella 
from Malaya, much simplified, x 1000. 

with longer and longer, tapering internodes ending in acute colourless tips' 
0*5 jit wide or less. The dichophyses also become smaller toward the 
hymenium where they are usually coralloid clumps so dense and closely 
set that the basidia project between them at wide intervals and never build 
a continuous layer. In one species that I have seen the hymenial dicho- 
physes are shaped like the digitate cystidia on the oi Androsaceus 

(Marasmieae) and may even be simple, but such are merely rudimentary 
states, and true setae seem not to occur. Gloeocystidia may or may not be 

' 26-2 

244 Transactions British Mycological Society 

present. The hymenium may remain as a vague layer or it may thicken, 
either continuously by the outgrowth of new hyphae to form dichophyses 
and basidia or intermittently in seasonal layers. 

The xanthochroic species are related to Lachnocladium and I have found 
in Malaya a pleuropodal ‘Stereum’ with the same construction. The 


Fig. 9. Stages in the development of the dichophyses of a xanthochroic species 
of Asterostromella from M2i\2iy^.y x 1000, 

cntical genus seems to be Hymenochaete, the tropical species of which are 
certainly of diverse afiinity, some having the dichophyses of Asterostromella, 
others the skeletal hyphae oi AsUrodon. The genus may provide instances of 
transitions between dichophyses and stellate setae, which appear to be 
regulanzed dichophyses with simple arms. I have not seen any xantho- 
chroic Polyporoid or Hydnoid fungi with dichophyses. ^ 

Notes on Asterostroma and Asterostromella. E. J. H. Comer 245 


The structure oi Asterodon ferrugimsus is described. It is dimitic with stellate 
setae, extrahymenial setae and hymenial setae, as well as skeletal hyphae. 
The shape of the setae and the direction of the skeletal hyphae express the 
action of the form-factors of the fruit-body. The fungus seems to be most 
suitable for experimental investigation of these obscure processes. 

The structure of is further illustrated by reference to Astero- 

stroma and the xanthochroic species oi Asterostromella, Hymenochaete appears 
to be the critical genus for the elucidation of the structure and affinity of 
these fungi. 


Bourdot, H. & Galzin, A. (1928). Hymenomydtes de France. 

Corner, E. J. H. (1932 a). A Pomes with two systems of hyphae. Trans. Brit. myc. Soe. 

XVII, 51-81. 

Corner, E. J. H. (1932^). The fruit-body of Polystictas xanthopus. Arm. Bot.^ Lond.^ xcvi, 

Corner, E. J. H. (1932^). The identification of the Brown Root fungus. Gard. Bull. S.S. 

V, 317-50. ^ 

Donk, M. a. (1933). Revision der Niederlandischen Homobasidiomycetae-Aphyllo- 
phoraceae, II. Med. Bot. Mus. Herb. Univ. Utr. ix, 1-278. 

Patouillard, N. (1900). Essai taxonomique sur les families et les genres des Hymenorry cites. 
PiLAT, A. (1935). Additamenta ad Floram Sibiriae Mycologicam. Bull. Soc. mycol. Ft. 
LI, 414. 

Rogers, D. P. & Jackson, H, S. (1943). Synonymy of some North American Thele- 
phoraceae and other resupinates. Farl. i, 263-328. 

{Accepted for publication 24 February 1947) 

[ 246 ] 



By £• G. JEFFERYS, Imperial Chemical Industries Limited^ 

• Buttemich Research Laboratories^ The Fry the ^ Welwyn 


Most techniques previously described for the determination of the pro- 
duction of antifungal substances by fungi or other organisms have involved 
growth of the fungi on liquid media^ serial dilutions of the culture filtrate 
being then assayed. 

Weindling (1934) produced culture filtrates of Trichoderma lignorum and 
observed microscopically the effect of serial dilutions on young hyphae of 
Rhizoctonia solani. Brian and Hemming (1945) again used serial dilutions of 
culture filtrates. These solutions were tested to determine the highest 
dilution at which they prevented germination of the spores of the test 
organisms. Irving, Fontaine and Doolittle (1945), using solutions of an 
antifungal substance produced by tomatoes, employed an assay technique 
analogous to the cylinder plate technique used in penicillin assay, the 
activity being determined by the diameter of a zone of inhibition. 

These techniques are more in the nature of quantitative assays rather 
than preliminary sorting tests. 

The use of direct inhibition of the growth of one fungus by another in a 
streak test was shown by Brian, Curtis and Hemming (1946) to be possible, 
provided that the test fungus is a yeast-like, rapidly growing form. They 
used Endomycopsis albicans. Tlm type of test, widely used for demonstration 
of antibacterial substances, cannot be satisfactorily used with mycelial fungi 
as the test organism. 

Where a large number of fungi are to be tested for antifungal activity, 
the above tests (except the streak test which has other disadvantages noted 
later) are liable to become very cumbersome, and it was considered 
desirable to develop a rapid technique using as little time, labour and 
material as possible. After a number of experiments on various lines, the 
following technique was developed. 


A 7 mm, diameter glass rod is bent to form a fiat rectangular cell of i by 
2 in. internal dimensions. This cell is sterilized by flaming and immediately 
placed in a sterile Petri dish. A small quantity of molten, sterile Czapek- 
Dox agar is poured into the cell, the quantity being only sufficient to flow 
round the edge of the cell, sealing it to the base of the dish on cooling. 
Other agar media can of course be used for fungi which do not grow 
satisfactorily on Czapek-Dox, When the agar has set, the cell is completely 

Production of Antifungal Substances. E. G. Jefferys 247 

filled with agar and again left to cool. Some agar usually flows over the 
edge of the cell or through the gap left between the two ends of the rod and 
this is removed with a sterile scalpel or spatula. The upper agar surface is 
then inoculated with a streak of the fungus under test. After two to three 
days incubation at 25° C. (or when good growth has taken place) the cell 
is gently prised off the bottom of the dish with a scalpel, picked up with 
forceps, toned over and transferred to a second sterile Petri dish, sterile 
precautions being observed throughout. The newly exposed surface is then 
inoculated with a spore suspension of the test fungus {Botrytis Allii Munn 
being frequently used in these laboratories). After a further sixteen to 
eighteen hours incubation the upper surface is examined with the low 
power of the microscope to determine the degree of germination. 


In the following table, germination is classified on a 0-4 basis as follows: 

0 No germination 2 20-50 % germination 

(i) Up to 5 % germination 3 50-80 % germination 

1 5-20 % germination 4 80-100 % germination 

It should be noted that little significance is attached to the difference 
between class 3 germination and that in class 4. 

It is often possible to note a gradation of germination between the spores 
oi Botrytis directly over the streak of the fungus under test (column (i)) and 
those away from the streak (column (ii)). 

Germination of Botrytis AUii in glass cell activity trial on Czcpek-Dox 

Fungus tested 



Antibiotic produced 

Absidia glama Hagem 




Botrytis Alin Munn 




Fusarium caeruleum (Lib.) Sacc. 




Fusarium graminearum Schwabe 




Metarrhizium glutinosum S. Pope 




Pmicillium Gladioli McCull. & Thom 


Gladiolic acid 

Pmicillium Janczowskii ZaL 



, ‘Curling-factor’ 

Stachybotrys atra Corda 



Not yet isolated 

Stemphylium sp. . 




Stereum purpureum Pers. 




Thamnidium elegans Link 




Trichoderma viride Pers. ex Fries 


Trichoderma viride Pers. ex Fries 

— , 



Uninoculated controls (2) 




It will be seen that germination of the spores of Botrytis after sixteen 
hours incubation is always suppressed or absent where an antibiotic is 
known to be produced. The results are consistently repeatable. 


This technique has several advantages over those previously reported. It is 
quick, taking only about four days to complete. The glass cells are simple 
to make and easy to handle though it should be noted that care must be 

248 Transactions British Mycologiccd Society 

taken when removing the agar-filled cell to insert the scalpel under the 
agar as well as the glass in order that both are transferred together. This 
procras is quite easy after a little practice and is facilitated by having cells 
no wider than i in. Finally the technique is very useful with quickly 
growing fungi which, in the streak test, are very liable to overgrow the 
test organism before any result can be obtained. 

Although this technique has been used only for demonstrating the 
production by fungi of antifungal substances, it could probably equally 
well be used to inyestigate the production by bacteria and actinomycetes 
ot substances antagonistic to fungi, bacteria or actinomycetes. 


A new technique is described for rapidly determining whether a mould or 
other micro-orgamsm produces antifungal metabolic products. This requires 
the use of a small rectangular glass cell filled with agar of which one side 
IS inoculated _wth the fungus under investigation and the other side is later 
inoculated with a standard test fungus. 

ad^ce valuable help and 


Brian P W., Curts, P. J. & ^mming, H. G. (1946). A substance causing abnormal 

S; 2.1. Tr®, w! 

Irving, G.W., Fontaine, T. D. & Doolittle, S. P. (1045), Lvcooersicin a f 

agent from the tomato plant. Science, cii (2636), 9-1 l ^ ^ fungistatic 

rS ^ in the parasitic action of 

I ^^gnorum on Rkzoctoma solam and other fungi. Phytopathology, xxrv (i i),^ 

{Accepted foT publicdtion 20 Februdry 1947) 

[ 249 ] 


By H. F. DOVASTON, West of Scotland Agricultural College 
(With Pl^te XIV and 7 Text-figures) 

Diedicke (1903) mentioned a Helminthosporium leaf-spot of perennial rye- 
grass in Germany. He remarked that lesions were produced resembling 
those caused by H. teres on barley and H. Bromi on Bromus inermis (i.e. they 
were net-lesions), but he found no associated ascigerous stage and published 
no further details. Drechsler (1923) described from the United States a 
Helminthosporium leaf-spot of Lolium mult^rum Lam. (X. italicum A.Br.) and 
of L. perenne L. under the name of Helminthosporium siccans n.sp,, and this 
organism was said to produce only spot lesions on these hosts. In the same 
paper he described a closely related species, H. dictyoides, which formed 
ntt-lesions on Festuca elatiorij. {F. pratensisHaAs.). 

It has recently been pointed out that the study of graminicolous Helmin- 
thosporia has been neglected in Britain (Dennis & Wakefield, 1946), and 
the first British record of ryegrass leaf-spot was published by Sampson and 
Western (1940), although the disease had been familiar to Miss Sampson at 
Aberystwyth since 1922. Material was collected in Scotland by Dennis in 
1932, but the record was not published until ten years later (Dennis & 
Foister, 1942). There is little doubt that the disease has been overlooked, as 
it is relatively common in Scotland and may be found on wild populations 
of Lolium perenne remote from cultivation. 

The types of lesions reported on the two species of ryegrass are sum- 
marized in the following table : 

Table i 

Author Country 

Diedicke, 1903 Germany 

Drechsler, 1923 U.S.A. 

Sampson and Western, 1940 Wales 

Dennis and F oister, 1 942 Scotland 

Lesion type 

Apparently net lesions 
Spot lesions only 
Spot lesions only 
Net and spot lesions 

There can be no doubt that in Scotland both spot and net lesions are 
present on both perennial and Italian ryegrass, but in general net lesions 
are most clearly developed on Italian ryegrass. 

Sampson and Western (1940) concluded that their material corre- 
sponded to Helminthosporium siccans DrechsL, but the nature of the lesion 
types and the specific identity of the British organism or organisms will 
be described in a later paper. It is proposed to describe here only the 
ascigerous stage of a Helminthosporium which causes net lesions on Italian 

During the last two years large numbers of monospore cultures have been 





00 . 



Pyrenophora/rom Italian Ryegrass. H. F. Domston 251 

were then picked off under a binocular microscope with a dry needle and 
transferred to oatmeal agar. 

Cultural characters are similar to those described by Sampson and 
Western (1940): light grey aerial mycelium, olive-green plectenchyma, 
and spore production abundant after three weeks. Development of 
sclerotia is variable in different isolates : sometimes very few are present, 
but occasionally they are numerous and large. Generally speaking, when 
sclerotia are well represented development of aerial mycelium and 
Helminthosporium spores is correspondingly sparse. The sclerotia are filled 
with an oily filamentous tissue of vertical orientation but indefinite 
structure (PL XIV, fig. 4), and externally they bear flexuous tapering setae. 
They are often well developed in^ cultures made bn sterilized ryegrass 
straw, but until recently only traces of asci have been observed. However, 
one culture made on oatmeal agar in September 1946 produced in 
February 1947 a quantity of ripe asci and spores, which proved that the 
sclerotia were immature perithecia and confirmed the general prediction of 
Drechsler. Commenting on the appearance of subspherical sclerotia in 
cultured material of several species he remarked : ‘ The writer is inclined to 
believe that in whatever species such sclerotia or abundant anastomoses 
(resulting in the production of complexes of lobulate segments) are found 
to occur, perithecia may be sought with considerable prospects of success.’ 
The anastomoses referred to are abundant in the plectenchymatic layer of 
the ryegrass Helminthosporium. These cultures made in September 1946 were 
kept outdoors, and the very cold winter of 1946-7 may have facilitated the 
production of asci and spores, as Drechsler found with Pyrenophora teres that 
‘ abundant moisture and a relatively high temperature appear to favour 
the conidial stage, and it is not illogical to expect that a long protracted 
period of cold, dry weather in Spring might result in the production of 
more nearly normal spores’. Temperatures were very low at Auchincruive 
during January and February 1947, and more than once reached 0® F. — a 
very rare condition in the west of Scotland. 

Pyrenophora Lolii Dovaston sp.nov. 

Perithecia inaequaliter conformata, 300-1 000 ju diametro (av. 710 x 
550/>t.), maturitate rostello conico brevi praedita, 1-50 setulis flexuosis 
multiseptatis attenuatis ad 200//- longis basi 8~iija crassis omata. Asci 
octospori, numerosi, 172-155-230 x 27*-^4‘”4i/x, late cylindrici, apice 
truncati, annulato-incrassati, basi pedicillati, pedicello quam in P. Avenae 

P. tereti lon^ovc. Sporae pallide brunneae, 49-55-67 x i6-i5-22/i, 
plerumque transverse 5-septatae, longitudinaliter o-4-(saepe 2-) septatae. 
Sporae saepe abortivae, sed plurimi asci 2-6 sporas perfectas exhibent. 
Germinatio celeriter evenit, tubulis germinationis lateralibus 1-5 (ple- 

Hab. in substrate solido artificiali (^oatmeal agar’ appellato) culta: 
culturae e Helmint^^ sp. maculas reticulatas in 

italico Huds. excitantis oriundae, Auchincruive, Ayrshire, Scotia. 


Transactions British Mycological Society 

Pyrenopkora Lolii Dovaston n.sp. 

Perithecia irregular in shape, 300-1000/4 in diameter (average 
710x550/4), with a short conical beak when mature, externally 1-50 
flexuous, tapering, multicellular setae may be present 8-11/4 in diameter 
at the base, and up to 200/4 long. Asci 8-spored, numerous, 1 72-1%- 
230x27-^^-41/4 broadly cylindrical when mature, apex truncate and 
with a thickened apical ring, stipe longer than in P. Avenae and P teres 
Spores pale brown, 49-5(9-67 x i6-/.?-22/4 nearly always with 5 trans- 
verse septa when mature and with o to 4 (mostly 2) longitudinal septa 
Spores frequently abortive, and most asci show 2 to 6 normal spores 
Germination prompt, with i to 5 (mostly 2) lateral or polar germ tubes' 

Habitat. In a single spore culture on oatmeal agar from a Helminthosporium 
which produced net lesions on Lolium italicum Huds., Auchinc^ve 
Ayrshire, Scotland. ’ 

Conidiophores and spores of a Helminthosporium often develop on the 
upper surface of the perithecia, and these correspond in morphological 
characters with H. siccans Drechsl., but they have not yet been compared 
with American material. ^ 

^ A comparison of the above measurements with the corresponding ones 
for the related Pyrenopkora teres Drechsl., P. Bromi Drechsl., and P. Avenae 
Ito and Kuribay, shows that the present organism more closely resembles 

P. Avenae thdoci do the oth^x two, 

Appro^mate modal measurements in Pyrenopkora species are given in 
Table 2. The data for P. teres and for P. Bromi are from Drechsler^(i923) 
and for P. from Dennis (1934). It should be borne in mind that £ 
gures for P. Zo& are based on a single collection, and that the material of 
P. Avenae and P. teres was limited. 

Table 2 

P. Bromi 

P. teres 

P. Avenae 

P, Lolii 

Diameter of perithecia 
Size of asci 

Size of spores 

No. of transverse septa 


















Ratio: length/width of 



(mostly 5) 



and^/lv ^ eccentricity of the spores, 

snores' o?P ellipsoidal of the four. The 5-septate 

Tlveme^n f^^inguish this species from P. teres, and it differs from 

£tac X smSer ^ perithecium, the rather broader 

D ’ narrowly eUipsoidal spores, 

four months still pathogenic to Lolium italicum after 

and produced indefinite net lesions four to five days 
als^maS b^fr^/hT'^^ of two to three spores only.) Attempts were 

of seed which was th ^ oats by leaf inoculation, and by inoculation 

ot seed which was then germinated m stiver sand. Barley was not attacked, 

Pyrenophora/rowj Italian Ryegrass. H. F. Dovaston 253 

but there were indications of slight pathogenicity to oats. This will be dealt 

with in a later paper. 

I am indebted to Miss E, M. Wakefield who kindly prepared the Latin 
diagnosis, and to Dr R. W. G. Dennis for several useful suggestions. 


Dennis, R. W. G. (i 934 )‘ Notes on the occurrence of Pyrenophora Avenae I to & Kuribay, 
in Scotland. Trans. Brit. myc. Soc. xix, 288. 

Dennis, R. W. G. & Foister, C. E. ( 1942). List of diseases of economic plants recorded in 
Scotland. Trans. Brit myc. Soc. xxv, 274. 

Dennis, R. W. G. & Wakefield, E. M. (1946). New or interesting British Fungi. Trans. 
Brit. myc. Soc. xxix, 159. 

Diedigke, H. (1903). Ueber den Zusammenhang zwischen Pleospora- und Helmintho- 
sporium-Arten. ^bl. Bakt. II, ix, 317. 

Drechsler, C. (1923). Some graminicolous species of Helminthosporium. I. J. agric. Res. 

XXIV, 641. 

Sampson, K. & Western, J. H. (1940). Two diseases of grasses caused by species of 
Helminthosporium not previously recorded in Britain. Trans. Brit. myc. Soc. xxiv, 255. 

Figs. 1-6. Pyrenophora Lolii n.sp. 

Fig. I . Three abortive spores and one normal spore in an ascus, living, x 600. 

Fig. 2. Ascus showing remains of five abortive spores, living, x 500. 

Fig. 3. Ascus containing eight normal spores (six are visible at this focus), living, x 650. 

Fig. 4. Indefinite granular tissue from a squashed perithecium, one normal and two abortive 
spores are visible, living, x 300. 

Fig. 5. Living ascus after twenty-four hours in water, direct germination has taken place of the 
unorganized ascus contents through the ascus wall, x 300. 

Fig. 6. Net lesion on Lolium itdicum, from the plot which produced Pyrenophora Lolii, living, x 3. 

Fig. 7, Helminthosporium spore from the culture which also produced P. Lolii. Lactophenol 
preparation, x 700, 

{Accepted for publication 2 April 1947) 

[ 254 ] 



By H. V. NE'WBY, Department of Botar^, 

Queen Mary College, University of London 

(With 23 Text-figures) 

Saprolegnia anisospora was described in 1888 by de Bary, who had collected 
it in the neighbourhood of Strassburg. The species has remained unique 
among its family from a combination of two outstanding characteristics: 
(^) the production of giant zoospores (in addition to others of smaller 
sizes) ; and {b) the excentric structure of the mature oospores. The latter- 
feature was held in doubt by Coker (1923)5 who described two forms 
collected in North Carolina as belonging to this species. Coker described 
these forms as possessing centric oospores ; but in other important characters 
they appeared to agree closely with de Bary’s description. This close 
resemblance covered a distinctive combination of characters and Coker 
wrote: ‘One is inclined to suspect that de Bary, who rarely made a mistake, 
was in this case wrong in thinking the normal eggs excentric. His figures 
clearly show excentric eggs, but may they not have been breaking down? 
This seems the more likely as no other Saprolegnia has an excentric egg.’ 

I was fortunate enough to isolate this interesting, but little-known, 
species, and observations were made upon bacteria-free, single-spore 
cultures over several months. The species was found to agree in detail with 
de Bary’s description, in so far as comparison can be made without re- 
corded measurements of the original form; for de Bary’s otherwise careful 
description included no measurements of the reproductive organs. In 
particular, it has been possible to confirm the character held in doubt by 
Coker, namely, the production of oospores having a typical excentric 
structure. The species has been reported in Latvia by Apinis (1929), who 
also found excentric oospores. I have found no previous record of the 
occurrence of this species in Great Britain. Many differences are apparent 
between the present form and the two American forms described by Coker, 
and the question of a different oospore structure in the American forms 
calls for consideration. The following account includes essential measure- 
ments made under stated conditions of culture; the form and behaviour of 
the zoospores is described in some detail; and the character of oospore 
structure is discussed. 

The fungus was isolated from a dead roach {Leuciscus rutilus)^ collected 
near Haslemere, Surrey. The material was first taken from a patch of 
felted hyphae on one side of the fish, and a bacteria-free stock culture 
prepared by growing the fungus upon an agar (Barnes’) medium, and 
removing inoculum from the under-surface of the agar. The sub-cultures 

Observations on Saprolegniaceae. H. V. Newby 255 

thus obtained were tested by inoculating a liquid maltose-peptone medium 
(which rapidly became turbid in the event of any persisting bacterial con- 
tamination). This medium, which had the composition: maltose 5-0%, 
peptone (Parke Davis) o*i %, proved useful for maintaining stock cultures 
over periods of several months without sub-culturing. Single-spore 
cultures were prepared by using a dummy microscope objective with a 
cutting tube of small bore, enabling any selected spore to be cut out, 
together with a small block of agar, from an agar plate. The latter was 
prepared a few hours previously by 'streaking’ the surface with a suspension 
of encysted zoospores, using a platinum loop of triangular shape. 3 % agar 
yields a firm gel which does not stick inside the cutting tube. 

The following description, except where otherwise stated, relates to 
observations upon single-spore cultures grown on sterilized hemp seed, in 
approximately 45 c.c. of sterilized glass (Pyrex) distilled water, within the 
temperature range 12-16'' C. Comparative cultures, in which segments of 
sterilized mealworm, approximately 1-5 mm. long, were used in place of 
hemp seed, showed no important differences. Not more than two seeds (or 
mealworm segments) were used in any one culture vessel. A glass dish, 
measuring 6 cm. in diameter by 3*5 cm. in depth, with drop-on lid, was the 
type generally used. 

Habit and general course of development 

It is felt that this aspect has often received insufficient attention in descrip- 
tions of members of Saprolegniaceae, many of which show distinctive 
features in their habit and general course of development when grown 
upon suitable baits in water-culture. 

Under the conditions stated, this species developed a main turf which 
appeared delicate but tense, relatively short, reaching a length of approxi- 
mately 4-5 mm. in five to six days and exceeding this only in old cultures 
of four weeks’ growth. Zoosporangia were numerous after three days. 
Oogonium initials became abundant in five to six days, forming a dense, 
opaque zone in the outer region of the main turf. In old cultures the limit 
of the main turf remained well marked by this dense zone, although a 
delicate outer turf, bearing some gemmae and markedly smaller oogonia, 
was often present. At a constant temperature of 10° G., the main turf 
usually reached 5-7 mm. in length and zoosporangia were more abundant; 
at temperatures between 17 and 20° G., the turf rarely exceeded 4 mm. 
and few zoosporangia were produced. 

A. Z^osporangia 

Primary zoosporangia varied from almost cylindrical to a prevalent type 
which was broadest either above or below the median point and often 
slightly bent or asymmetrical about the long axis (Figs. 5-8) . The majority 
were between 130 and in length and 24-26^6 in breadth, which 
exceeds the dimensions given by Goker. A well-marked, button-like 

Observations on Saprolegniaceae. H. V. Newby 257 

papilla at the tip differed from the usual Saprolegnia pattern in being com- 
paratively broad, slightly funnel-shaped, and almost flat across the top, 
where the wall was markedly thin. This contrasted with the remainder of 
the papilla which had a wall somewhat thicker than that of the sporangium 
itself (Figs. 8 and 13). Proliferation occurred repeatedly and the first 
few secondary sporangia were often confined within the empty primary 
sporangium; later sporangia in the same series commonly showed partial 
emergence, the protruding part acquiring a characteristic vase-like form 
and showing a marked tendency, when empty, to become telescoped back 
upon the preceding sporangium (Figs. 9-1 1). Long series of proliferated 
sporangia were invariably found after a few days’ growth and were 
specially evident in cultures grown at 10° G. Secondary sporangia formed 
by lateral renewal (as in Achlya) were comparatively rare. 

B. Zoospores 

Observations on the present form confirm de Bary’s report that all the 
zoospores from a given sporangium ate of the same order of size, and 
further, that this bears no apparent relation to the size of the sporangium. 
The zoospores from a given sporangium tend to fall within one of three 
size-groups : giant, intermediate, or . small. In the present form the 
number of zoospores in any sporangium was never large, commonly 
between ten and twenty-five, rarely as few as three or four. 

(a) First motile stage 

Measurements were made, as rapidly as possible, of spores newly 
liberated from sporangia. Although such measurements must necessarily 
be approximate, they have been as far as possible confirmed by subsequent 
measurements of spores killed with iodine, when, however, a slight con- 
traction occurred. 

Table I. Dimensions of zoospores 


ist motile stage: Len^ 

Maximum width 

Encysted zoospores 
I St stage: Diameter 




( m ) 











Legends to Figures i-x I 

Fig. i. Two youhg oogonia with antheridia associated and with a dilation in the parent hypha. 

Fig. 2. Yoimg oogonia with diclinous antheridia. 

* Fig." 3. Young oogonium, developed immediately beyond an old zoosporangium, with diclinous 
antheridia associated. 

Fig. 4. An oogonium with mature oospores and with empty antheridia still attached.^ The 
oospores show the characteristic exeentric structure and the thick two-layered wall, the inner 
layer showing stratification. 

Figs. 5-7. Primary zoosporangia. 

Fig. 8. Primary zoosporangium containing spores pf intermediate type, immediately prior to 
discharge. The characteristic features of Ae apical papilla are well marked, * 

Figs. 9“i I. Zoosporangia in various stages of proliferation. Fig. 9 shows the characteristic vase- 
like form of the emergent part. Figs. 10 and irshow the ‘telescoping’ back of the emergent 

■: •■■■parts. ■ ■ 



Observations on Saprolegniaceae. H. V. Newby 259 

Cysts of less than 13*5^^. diameter were rarely found and it appears 
probable that spores of both intermediate and small types, when encysted, 
all belong to the diameter range of iS'S-iS'S/x. It will be seen that the 
dimensions of the zoospores fit de Bary’s description: 'The largest of these 
(zoospores) are more than double the size of the smallest, though there are 
also intermediate forms/ De Bary made no mention of the relative 
frequency of the smallest spores : in the present case they have been less 
numerous than the other two types. He described the large spores as 
' almost equalling that (the size) of the oospores and having very dense, 
granular protoplasm’ ; with which the present form exactly agrees. 

The following notes refer to giant and intermediate spores. These 
showed considerable variety of form in their first motile stage. A large 
number showed a broadly pointed anterior end and were approximately 
rod-shaped, or slightly narrower about the middle. Elongate-pyriform and 
pyramidal forms were also frequent and many of these had their anterior 
end drawn out into a definite beak. Any of these types occasionally showed 
two short lobes at the posterior end (Fig. 12). In old cultures, grown at a 
constant temperature of lo"^ C., some unusually broad, giant spores have 
occasionally been observed, measuring approximately 27/t in length by 
2011 wide. This recalls Coker’s description : ' In nearly all cultures there are 
formed, in addition, a few very large spores at least twice the bulk of the 
ordinary large spores’; although in the present case these have not been 
observed to occur as frequently as his statement would indicate. Occasional 
double spores have been observed: these have two sets of flagella and 
clearly result from an incomplete cleavage of the protoplasm in the 
developing sporangium, Coker has figured such compound spores for this 
species (Coker, 1923; plate 7, fig. i). 

The main body of a zoospore, composed of dense, coarsely granular 
protoplasm, contrasted strikingly with the usual, delicately granular 
appearance of zoospores of the Saprolegniaceae. A distinct hyaline area 

Legends to Figures 12-23 

Fig. 12 A-F. Giant zoospores. Flagella not drawn. Vacuoles are seen in A, G, E and F. 

Fig. 13. Sporangium containing giant spores, just before discharge. The hyaline anterior ends of 
the spores are seen. Several spores showed active amoeboid movements. 

Fig. i 4A~D. Stages in the escape of a zoospore from its primary cyst. 

Fig. i 5A-~C, Changes in shape of the emerged zoospore (second motile phase), immediately 
following the events shown in Fig. 14 A-D. 

Fig. 16 A, B, Two views of a zoospore during the second (reniform) motile phase, showing the 
two flagella in Fig. 16 A and the lateral groove from which they arise (Fig. 16B). 

Fig. 17. A zoospore of the first motile phase and of intermediate type, trapped at the opening of 
the sporangium. The two apical flagella are shown. 

Fig. 18. Excentric oospore with thick, two 4 ayered wall and a single, large fat droplet. The 
protoplasmic contents are limited to a thin, cup-like layer. 

Fig. 19. Excentric oospore having a second, much smaller droplet. 

Figs. 20 and 21. Immature oospores containing several droplets and with wall still relatively 
■ .'thin. 

Figs. 22 and 23. Oospores with characteristic thick wall and apparently near maturity, but 
containing a few separate fat droplets. Developed at approx. 20° C 


26 o Transactions British Mycological Society 

Sf f ported anterior end, from which a pair of lon^ 

flagdla arise. The flagella could sometimes be distinguished momentarilf 
on the Imng spores, but they became more distinct on spores killed with 

i^ine, when they were seen to exceed the length of the spore body fFiff 17? 

1 ^ continuous vibratory motion, apparentlv 

related to the movements of the flagella, and also side-to-side m??ementl 
m a rapid, sensitive manner. The remainder of the spore body commodv 
showed some ca,paaty for amoeboid change of shape and this was frJ 
quently observed even before &e spores escfped from the sporangium 
the mam body of the spore a few minute, rounded, refractive bodies were 
evidence, the most co^picuous of these was often close within the 
pen;^ery of Ae spore, shghtly in advance of the middle fFiff 12 A C 
and F) No alteration in the size of this refractive spot could be detected 
dunng the bnef ghmpses ofiered by the living spores aetected 

The giant zoospores swim comparatively slowly during dieir first motile 
stage, revolving at the saine time slowly upon their long axis. They were 
observed to remain in motion for varying periods of five to thirty mfnutes 
They frequently showed an interesting behaviour when coming to rest the 
spore was seen to stop, turn itself round shortly, a few timfs after Ihl 
m^ner of a dog preparing to lie down, and then finally contract into a 
sphere. Shordy afterwards, a thin cyst wall became discLnible 
The small type of zoospore is pyriform and has less dense protoplasm. 

(b) Spore discharge 

Ir^ediately prior to spore discharge, observations on giant and inter 
mediate spores showed distinct amoeboid movements some nf Ae o * 
beco^ng much elongated in attempting ^ 

S?of sporangium. At this stage the hyaline antSS 

cncls of tlic spores were usuallv clearlv , 

were usually arranged in single file. ^ ^ o or three spores 

The^ topmost spore was frequendy placed with its antermr er,^ • *• 

away from the tip of the stioraneinm ^Tn its^ anterior end pointing 

JLe of the »d giX 


nunute. t' r""" 


Observations on Saprolegniaceae. H. V. Nevohy 261 

spores emerged in a steady succession, not explosively. As each spore made 
its way through the open papiUa it became somewhat constricted, and 
often remained somewhat dumb-bell shaped for a few seconds after release. 
After the first few spores had escaped, the later ones moved up to the 
opening more slowly. As far as could be discerned, a spore showed no 
connexion with others during spore discharge; and the last spores to 
emerge were often widely separated as they passed slowly up to the 

(<;) Second motile stage 

The escape of the second motile stage from cysts measuring approxi- 
mately 1 8*5^ in diameter was observed (Figs. 14 and 15). It was usually 
accomplished in one to two minutes ; but in some instances the spore, 
having emerged, then remained just outside the cyst wall for a further 
period of as much as thirty minutes, before beginning to swim : during this 
period it was almost stationary, except for slight rocking movements. (It is 
possible that such delay before beginning to swim may have been caused by 
the light or temperature conditions upon the microscope stage.) In the 
second motile form the spores showed the same coarsely granular appearance 
of the protoplasm. They were ovoid, with slightly pointed ends and a lateral 
groove running back from the anterior end almost the entire length of the 
spore body. A median pair of long flagella arose from this groove (Fig. 16). 
Typical dimensions were as follows: a larger type, length i8*5~20*5fx, 
width i2-i3*5/x; a smaller type, length 15/i, width i2jL6. The spores swim 
much more rapidly than in the first stage, also revolving quickly upon their 
long axis. Spores emerging from some cysts which had become fortuitously 
imprisoned inside an empty sporangium were observed to swim per- 
sistently backwards and forwards, exploring repeatedly the inner surface of 
the sporangium wall, and showing some capacity for momentary change of 
shape when reversing direction. The spores have been observed to remain 
actively swimming for upwards of thirty minutes, or in other cases to come 
to rest relatively soon. After rounding off*, each spore formed a spherical 
thin- walled cyst of approximately the same size as that from which it 

In most cultures, a small proportion of cysts measuring only 
has been observed. In some cultures, three to four weeks old, which had. 
been kept at a temperature of 10° C., such small cysts were relatively 
numerous and had settled in large groups upon some of the older hyphae. 
They appeared to have relatively scanty protoplasmic contents and some 
showed a slender, short germ-tube. It was not possible to determine if 
these small cysts were produced as a consequence of a polyplanetic 
behaviour under the conditions mentioned. 

{d) Spore size in relation to single-spore cultures 

De Bary made single-spore cultures from giant spores and reported that 
these cultures gave rise to plants which showed the normal behaviour in 
producing zoospores of various sizes. A long interruption in the present 
work resulted in the loss of the stock cultures and prevented this point from 

262 Trama£ti(m Bntish My cological Society 

being confirmed. Once only, a culture used for detailed observation had 
been made from an encysted spore of known diameter. The cyst diameter 
was 13/* and the cyst may therefore be taken as belonging to a spore of the 
small type. This gave rise to a plant which made a completely normal 
development and produced giant spores as well as intermediate and small 
ones. This observation is thus complementary to de Bary’s report. 

A. Oogonia and antheridia 

Oogonia were mostly spherical or sub-spherical, having a short neck or 
almost sessile, and appearing very early. The first oogonia were mainly 
terminal, but by the sixth day others were appearing on lateral branches 
and slightly later on lateral branch systems of cymose type. They formed 
a dense zone in the outer region of the main turf. The majority fell within the 
diameter range, 40-52JU., and contained two to six, or more, ova, commonly 
four or five. The oogonium wall was smooth, moderately thin, without 
visible pits (Figs. 1-4). A dilation of the hypha, some short distance below 
a terminal oogonium, or at a point of forking below two terminally borne 
oogonia, was frequently observed, and appears to be fairly characteristic 
of the species (Fig. i). 

Antheridia were diclinous and developed early on long, relatively 
slender hyphae, which later disappeared leaving the empty antheridia 
clearly distinguishable; at least two, frequently several, antheridia were 
associated with each oogonium, becoming closely wrapped about it, often 
to a large extent hiding the oogonium. The antheridia were commonly 
broad or appeared somewhat inflated, frequently lobed (Figs, i, 3 and 4). 
Two or three broad, cushion-hke ‘feet’ were sometimes visible where the 
antheridium was applied to the oogonial wall. The antheridial hypha often 
continued its growth, thus reaching a second or subsequent oogonia. An 
antheridial (fertihzation) tube was observed in a few instances. 

B. Oospores 

The oospores at maturity varied from 16 to 24/z. in diameter, most being 
1 8-2 1 /X. Apart from the excentric structure, the markedly thickened wall 
is another outstanding feature. In mature oospores the wall was 3-3*5 jx 
in thickness and distinctly two-layered: the refractive outer layer was 
approximately one-sixth of the total thickness (or a greater proportion 
in some instances which were apparently immature). The inner layer 
appeared denser, less refractive, and usually showed a faint, concentric 
stratification (Figs. 4, 18 and 19) . 

In most matinre oospores the contents showed a typical excentric 
structure, essentially sinular to. that which is characteristic of numerous 
spedes of the single lateral oil droplet was somewhat elliptical and 

the protoplasm of exceedingly fine, granular consistency (Fig. 4). In some 
instances, however, the oil droplet was much larger, occupying the greater 
part of the interior and limiting the protoplasm to a relatively narrow 

Obsewatiom on Saprolegniaceae. H. V. Newby 263 

crescent as viewed in optical section (Fig. 18). Other conditions were also 
observed, in which two or more smaller oil droplets were present instead of 
a single larger droplet, but these were frequently grouped to one side of the 
protoplasm, i.e. upon an excentric plan (Fig. 20). Since the wall of these 
multi-droplet oospores was relatively thin, it is suggested that they repre- 
sented immature stages, in which the several oil droplets had not yet 

The occurrence in old cultures, however, of oospores, with a thick, two- 
layered wall, and containing several oil droplets, often of varied sizes, 
indicates that a proportion of the oospores may never acquire a typical 
excentric structure (Figs. 22 and 23). Here it may be recalled that species 
of Achlya which typically possess excentric oospores, may at times show 
other types having two or more smaller droplets (Coker, 1923; plate 35, 
fig- 5? pl2.te 49, fig. 12). In the present species, at temperatures ranging 
from 17 to 22° C., oospores of the typical single-droplet, excentric pattern 
have been noticeably less frequent. Under these conditions, the matura- 
tion of the oospores becomes more rapid, the oospore wall quickly thickens 
to the maximum observed, and the several oil droplets present may show 
no indication of coalescence. It therefore appears that a more gradual 
development is most favourable for excentric structure. 

Oospores of the true excentric type have developed equally well in 
hemp-seed and mealworm cultures, also in two liquid media which were 
tried, viz. ; 

Solution I . Maltose 0*0125 % plus Witte’s peptone 0*025 %. 

Solution 2. Maltose 0*025 % Witte’s peptone 0*025 %* 


Regarding the unique character of this species de Bary wrote: Tt is 
distinguished from all others by the structure of the ripe oospores, which 
resemble those of Achlya polyandra. Along with this, there is the altogether 
peculiar feature of two kinds of sporangia, the one like those of allied 
species, the other forming giant zoospores. The largest spores are more than 
double the size of the smallest, but there are also intermediate types. The 
spores are all of one size in any one sporangium. There is no relationship 
between the size of a sporangium and that of its spores: sporangia of equal 
size will form spores of quite different dimensions.’ The present investigation 
serves to re-establish the altogether singular position of this species. The 
two forms described by Coker show many points of resemblance to de 
Bary’s species. The one apparent discrepancy lies in the structure of the 
oospores, which Coker has described as centric. His figures (Coker, 1923, 
plate 9, fig. 4), however, fail to convince me upon this point: thus, along 
with two apparently centric oospores, there is shown in the same oogonium 
an oospore which has eight conspicuous droplets grouped on one side of 
the protoplasm. This latter instance is not unlike immature stages which 
have been observed in the present form. It may be noted that none of the 
oospores figured by Coker shows the markedly thick wall which develops in 
the mature oospores of the present form, nor are they as thick-walled as 
those drawn by de Bary. From this it may perhaps be inferred that, in the 

264 Transactions British Mycological Society 

American, forms, the oospores were slow to mature, and that those studied 
by Coker were still relatively immature. 

There are other differences in the American forms which are summarized 
in Table 2: 

Table 2 

Zoospores : diameter of Diameter 

I St encysted stage of oospores 

American forms 
(according to Coker) 
Present form 


Intermediate Small 




Ip) Ip) 



13 - 7 -I 4-8 

1 0-5-1 1 *5 8-9 




1 5 * 5 - 1 3*5 



Coker described Ms forms as developing secondary sporangia by lateral 
renewal (as in Achlya) when grown in distilled water, but in spring water 
proliferation was as usual in Sapmlegnia. The latter method has been found 
typical of the present form when grown in distilled water, prepared in a 
Pyrex glass still. Further, oogonia borne in chains have never been 
observed in the present form; they were figured by Coker, but no details 
were given of the medium on which they were developed. The oogonia of 
Coker’s Form B are larger than those of the present form, but those of his 
Form A show a closer agreement. 

These differences of size may, however, be regarded as of minor sig- 
nificance when set against the combination of characters in which these 
American forms show agreement with de Bary’s species (including the, 
present form), Coker summarized these characters: ‘The small oogonia 
with unpitted walls, the small number of eggs, the numerous and con- 
spicuous dichnous antheridia on each oogonium, and the very variable 
spores.’ Observations upon oospore-development in the present form 
strongly indicate that, if the mature oospore structure of Coker’s forms were 
better known, they would be found to coincide also in this character; and 
thus to possess both those characters which mark out Saprolegnia anisospora 
as unique in its family. 

Summary ■ 

A detailed study of a strain of Saprolegnia anisospora colltcteA at Haslemere, 
Surrey, showed that it agreed in detail with de Bary’s original description 
of the species. 

In particular, the formation of oospores having a typical excentric 
structure has been confirmed. In this character, and in the production of 
zoospores of at least three size-groups, stands unique in its 


Detailed observations on the zoospores are recorded. 

The apparent difference in the American forms, described by Coker, in 
relation to oospore structure, is discussed. 

The main characteristics, as observed in hemp-seed cultures, grown in 
45 c.c. of glass (Pyrex) distilled water, at 12-16*^ C., may be summarized as 

Main turf 4-5 mm. Primary widest above or below the 

Observations, on Saprolegniaceae. H. F. Newby 265 

median point, often slightly curved or asymmetrical: majority i30- 
i95 X24“26/r. Terminal papilla broad, flat across the top, and having 
a thicker side wall. Proliferation occurs repeatedly, with partial emergence, 
empty protruding portions becoming telescoped back. Zoospores commonly 
10-25, and all of the same order of size, from any one sporangium: di- 
planetic. Three size groups were distinguished: giant, intermediate and 
small. Giant zoospores measured 29-32 x io-i2ft, their cysts 18*5-21 /a in 
diameter; their protoplasm is dense and coarsely granular and they show 
considerable amoeboid movement. Oogonia developed early and formed 
a dense zone in the outer region of the main turf, at first borne terminally, 
later in cymose, lateral groups: majority 40-52/^ in diameter, having 
usually 2-6 ova, wall not thick, unpitted. Antheridia diclinous, broad, 
several to each oogonium. Oospores typically excentric, having a thick, 
two-layered wall; majority 18-21 /a in diameter. 

In conclusion, the writer wishes to record his thanks to Prof. F. T. 
Brooks, F.R.S., and to Prof, F. E. Fritsch, F.R.S., for advice and criticism; 
also to Mr J. F. Leney, of Haslemere, who sent the specimen. 


Apinis, a. (1929). Untersuchungen iiber die in Lettland gefundenen Saproiegnieen. Acta 
Horti Bot. Univ, Latv. rv, 201-46. 

Bary, A. DE (1888). Species der Saproiegnieen. xlvi, 619-21. 

Coker, W. G. (1923). The Saprolegniaceae, Univ. N. Carolina Press. 

{Accepted for publication 27 May 1947) 

[ 266 ] 



By H. V. NEWBY, Department of Botany y 
Queen Mary College^ University of London 

(With 6 Text-figures) 

This species, described in 1899, was originally isolated from eggs of the 
sea-trout. Since Maurizio's original investigation it has remained com- 
paratively unknown. The only published record of its occurrence in Great 
Britain (Brown, 1938) makes no reference to those peculiar egg-containing 
structures, resembling antheridia or androgynous antheridial branches, 
which appear in this species. Maurizio furnished his account with only 
a single figure of these interesting structures. 

The present account records observations made upon single-spore 
cultures of this species. The fungus was isolated from a dead roach 
{Leuciscus rutilus) collected at Haslemere, Surrey, the material having been 
taken from a small patch of felted hyphae on the head of the fish. The 
fungus agreed well with Maurizio’s description and in the later stages of 
fruiting showed the anomalous egg-containing structures already men- 
tioned. Except where otherwise stated, the following notes refer to single- 
spore cultures, prepared firom bacteria-free stock culture, and grown on 
sterilized hemp seed in 45 c.c. of sterilized glass (Pyrex) distilled water, 
not more than two seeds being used in any one culture dish. The develop- 
ment of the fungus upon small segments (1-2 mm. long) of mealworm, but 
under otherwise identical conditions, was found to be essentially the same. 

General characteristics 

The main turf became well developed in six to seven days within a tem- 
perature range of 12-16^ G., reaching an average length of approximately 
7 mm. Later, a delicate outer zone develops, extending above the main 
turf to approximately i cm. above the hemp seed. In twelve to fourteen 
days after the beginning of the culture, a deep zone in which numerous 
oogonia and antheridia are developed becomes easily visible in the outer 
half of the main turf, and is conspicuous by the twenty-first day. A few 
spherical, gemma-like bodies, somewhat smaller than oogonium-initials, 
but with dense protoplasmic contents, are usually developed on the 
relatively slender hyphae of the outer zone; some are borne in series 
of two. 

The reproductive organs showed close agreement with Maurizio’s 
description upon most essential points. Intercalary oogonia, however, 
were comparatively rare, and none of the elongated, thread-like pattern, 
mentioned by Maurizio as containing up to seventeen ova in a row, was 

Ohsermtiom m Saprolegniaceae. H. V. Newby 267 

found. Oogonia were, for the most part, spherical or sub-spherical, with 
or without a short neck, and mostly borne on short, lateral branches. The 
maximum nurnber of ova observed was fifteen. Maurizio reported the 
number of ova in intercalary oogonia as, in some cases, approaching fifty; 
and Miss Brown (i 93 ^) figured oogonia containing considerable 
numbers of ova, from a strain collected in the neighbourhood of Aberyst- 
wyth. This strain was ascribed to Saprolegnia paradoxa^ although no 
antheridia were described. 

In the present form, every oogonium became associated with one or 
more, commonly three or four antheridia. These, as in Maurizio’s descrip- 
tion, were mainly of andro^nous origin. They were typically borne on 
short, curving branches, arising from the oogonial branch, or firom the 
main hypha close by (Figs. 2 and 4). Sometimes, however, diclinous 
antheridia, borne on relatively long hyphae, were present together with 
androgynous ones on the same oogonium. The prevailing type of anther- 
idium was club-shaped, with a well-marked angular bulge on the dorsal 
side. Often, an antheridium showed two or more lobes; and short, blunt, 
foot-like processes were commonly seen on the side applied to the oogonium 

Maurizio^s description indicates that the oospores showed a variable 
internal structure. In the present form, a well-marked sub-centric pattern 
was much in evidence, but others varied to an approximately centric type. 
The individual oil droplets appeared to be less distinct than is usual among 
Saprolegniaceae, even when viewed with a ^ inch oil-immersion objective. 
One curious double, or compound, oospore was found. 

Special variations of oogonia and antheridial branches 

In the later stages of fruiting, after two to three weeks’ growth, and more 
commonly, though not exclusively, at the higher temperatures employed 
(i7“"20° G.), a proportion of the oogonia was found to have anomalous 
structures associated with them. These usually had the form of one or two 
egg-containing outgrowths, arising either from the main body of the 
oogonium, close to the neck, or directly from the short neck itself. When 
relatively short, these outgrowths stand off stiffly firom the oogonium 
(Fig. 4), smaller ones often being somewhat narrowed at the base. The 
longer type usually arise from the oogonial neck, or immediately below the 
basal septum. They were frequently club-shaped, and curved upwards so 
as to bring the distal portion into contact with the oogonium wall. This 
type is therefore somewhat reminiscent of an androgynous antheridium 
(Fig. 3) , The number of ova in a single outgrowth varied between one and 
fever al; the ova were somewhat smaller than the average size within the 
oogonium proper, were arranged in a single row, and often laterally com- 
pressed to an elliptical form. The wall of an egg-containing outgrowth was 
thickened like that of an oogonium and pit areas were observed in some 
cases.' , 

A small proportion of oogonia were borne on short branches wHch were 
bent at an angle of almost 90° close beneath the oogonium, the point of the 

Fig. I. Young oogonium (u) with androgynous branches arising from its hypha One of these 

. ^ .-sin., 'em™! ifs 

^ Anantheridial (fertilization) tube is 

o^Dore^eStl^t ^ mtoal differentiation, also thickening oKhe wall, indicating 
oos^re aevelopment. The oogomum has a basal ingrowth. ^ 

beneaTh outgrowth which has arisen from immediately 

Pi*o- f ® anthendial branch is associated with the oogonium. 

Lnch®ISted*]^°sn^r°?-*^^ outgroij^s near the base and with one antheridial 
associated. An anthendial tube and a basal ingrowth are shown 

^ poiS°5°&^Se*^a f and having a short projection from the 

fiv fi nn^«!v: -fu ^ anthepdial tube is seen within the oogonium at (a). 

^ ala?eS associated, one of which is developing 

a lateral swelling which resembles an oogonium initial.^^^^^ ^^^ ^^^^^^^^^^ ^^ 

Observations on Saprolegniacem, H..V. Newby 269 

angle being frequently extended as a slight projection (Fig. 5). Sucli knee- 
like oogonial branches appear to be quite characteristic of this species and 
are mentioned by Maurizio. In the present form, the projecting knee-joint 
had, in some cases, all the appearance of a feeble attempt to form the type 
of anomalous outgrowth already described, although' in no case was such 
a projection large enough to have contained ova. 

The resemblances which some of the egg-containing outgrowths bear to 
androgynous antheridial branches have been mentioned. Other examples 
were observed in which an androgynous branch developed an egg-contain- 
ing structure in addition to one or more antheridia. In these, the egg- 
containing organ usually took the form of an asymmetrical swelling in 
a branch which was otherwise of average width. Such a swelling was com- 
parable in size to a small oogonium, had a similarly thickened and pitted 
wall, and^ contained usually not more than two or three ova. On the same 
branch, either above or below this structure, an antheridium was commonly 
developed, which became applied to the wall of the oogonium proper (that 
is, with which the androgynous branch was associated) (Figs, i and 6), In 
some cases, antheridia were seen to be applied to the egg-containing 
structure. itself, these antheridia arising on other androgynous branches, or 
on diclinous branches. In one of these cases an antheridial tube was 
clearly visible. The irregularly knotted character of antheridial branches, 
described by Maurizio, was not observed. 

An attempt was made to stimulate the production of these anomalous 
androgynous branches by growing the fungus in certain liquid media. 
Three of the solutions used (Nos. i to 3) are reported to have stimulated 
the development of antheridia and antheridial branches in other species 
(Klebs, 1899; Kauffman, 1908); solution 4 has yielded excellent crops of 
antheridia and oogonia with Saprolegnia parasitica and Sap, anisospora. The 
compositions of these solutions are as follows : 

Solution I. 
Solution 2. 
Solution 3. 
Solution 4, 

Leucin o*i % plus tri-potassium phosphate o-i % 
Leucin o-i % plus potassium nitrate o-i % 
Laevulose o*i % plus tri-potassium phosphate o*i % 
Maltose 0-025 % Peptone (Witte’s) 0-025 % 

None of these solutions had any effect in the direction required. They 
were, in fact, less successful in this respect than hemp-seed cultures. The 
lowest temperature range employed was 10-13^^ C., the highest 17-20° C. 
Abundant oogonia and antheridia were produced in solutions 3 and 4, the 
antheridia being in both cases mainly diclinous. In solution 3 occasional 
oogonium-like swellings on antheridial branches were observed ; in solution 
4 an occasional oogonium with a short protrusion was found, but no egg- 
containing outgrowths were present. In solution 4 the antheridia showed 
a strong tendency to be lobed or branched. Sometimes the antheridial 
branch forked immediately below the antheridia; in others, the branch 
remained simple, but the antheridium showed two, three or more relatively 
long lobes. The oogonia produced in solution 4 were practically in- 
distinguishable from normal oogonia grown on hemp seed; in solution 3, 
however, the oogonia showed a marked difference in that the majority had 

270 Transactions British Mycalogical Society 

a pronounced neck, frequently in length, in which one or more 

ova were tightly wedged. A few intercalary oogonia were developed 
in solution 4, but they were not of the elongated pattern described by 
Maurizio. Well-developed oospores were formed in both these solutions, 
although more tardily in solution 3. 


An account is given of some of the anomalous egg-containing structures 
observed during a study of this species, grown in single-spore culture on 
hemp seed. An attempt to stimulate their production by the use of 
synthetic liquid culture media is noted. 


Brown, E. M. (1938). Observations of Aquatic Fungi, Aberystwyth District. Trans, 
Brit rriyc, Soc. 'xxa, 

Kauffman, G. H. (1908). A Contribution to the Physiology of the Saprolegniaceae, Ann, 
Zorti., xxn, 361-87. 

Klebs, G. (1899). Zur Physiologic der Fortpflanzung einiger Piize (2). Jahrb,f, wiss, 
xxxiir, 513-93. 

Maurizio, A. (1899). Beitrage zur Biologie der Saprolegnieen. fur Fischerei, 

vn. Heft 2, 1^6. 

{Accepted for publication 27 1947) 

[ 271 ] 



(With Plates XV and XVI and 5 Text-figures) 


Three species of Myrothecium are dealt with here. Two of these, which are 
referred to in a previous paper (Preston, 1943), have not been previously- 
described and are considered to be new. The third, Myrothecium gramineum, 
was originally named by Libert (1837) on the printed label of a published 
exsiccatum. This species has been critically examined afresh and a fuller 
diagnosis of it is now presented. 

Myrothecium gramineumUib. 

(i) Material studied and characteristic features. 

Two collections of this species have been available. 

{a) An authentic Libert collection published in PL Crypt. Ard. exsic. 
No. 380 on decaying grasses in Herb. R. B. G. Kew. The original descrip- 
tion on this packet reads as follows: Myrothecium gramineum. Minutum 
rotundum in ambitu pilis erectis albis ciliatum; disco subturgido nigro; sporidiis 
cylindricis. In Gramineis putrescentibus. Aestate. 

(b) On Pennisetum subangustum; Makene, Sierra Leone; F. C. Deighton 
(M. 1743); 28. i. 1939 from Herb. Imp. Myc. Inst. No. 1494. 

The most characteristic feature of M. gramineum is the very stout and 
usually aseptate setae (see Text-fig. lA) which fringe the sporodochium. 
These make it readily distinguishable from the new species, next described, 
which has somewhat similar spores (see Text-fig. 2) but the setae of which 
are comparatively slender and always multiseptate. 

(ii) Revised description 

Mycelium (on potato-dextrose agar) . Pure white, forming a dense growth. 
Hyphae thin- walled, hyaline, septate; subaerial filament cells 20 X3-4/Z, 
aerial filament cells 20 x i~2ft. Sporodochia. Circular or elliptical, saucer- 
shaped, sessile, 0-2-0-9 mm. in diameter, o- 1-0*17 mm. deep. Black, 
surrounded by a fringe of numerous long, white, thick- walled setae. 

Straight, hyaline, tapering, ending in an acute or bluntly rounded 
point, generally continuous but occasionally with a single septum near the 
lower end about 40/X above the base; 200 to about 400/x in length, 
io-20jLc. wide near the rounded base and about 4/i just below the tip, walls 
3/4 thick at base tapering to about i/4 at tip, inner surface of wall often 
slightly sinuous, outer surface occasionally so. Conidiophores. of 

fertile hyphae and phialides . Fertile hyphae. Sub-hyaline, septate, oli- 
vaceous in the mass„ 2-3/4 wide with the terminal (or sub-phialide) cells 


Transactions British Mycological Society 

slightly broadening at the distal end; so closely intertwined as to be almost 
inseparable and forming a very compact sub-hymenial disk. Phialides 
Hyahne, straight or flexuose, slightly clavate or sub-cylindrical, 8-15 x 2«- 
generally arising in whorls of three or more from the terminal cells of the 
fertile hyphae; those arising from the lower filaments longer than those 
from the uppermost ones, the whole forming a dense palisade-like layer in 
which the individual phialides are very difficult to distinguish. Conidia 
Continuous, sub-hyaline or pale green, narrowly ovate, ends rounded or 
with the broader end occasionally slightly flattened, 7-10 x 2u averave 

8*5X2/i. ® 

Myrothecium jollymannii n.sp. 

(i) Material studied and characteristic features 

_ The type isolation of this species was originally made by F. W. Jollyman 
m 1936 from dried tobacco leaves in Nyasaland. A transfer was recced 
from the Imperial Mycological Institute as Jollyman 145, 14 ix 26 
Further transfers from this have provided all the material upon which’l^s 
description is based. 

The sporodochia bear a close superficial resemblance to those of M 
mundatum and of M. gramineum owing to the fringe of stiflf white setae which 
isxommon to each. The setae, however, are much more slender than those 
of gramtneurn and are multiseptate. They usually consist of seven to ten 
cells whereas those of M* inundatutriy which are even more slender rarely 
show more than five cells and the much more robust setae of M. gramineum 
^e eitoCT continuous or have but a single septum a short distance from the 
base The comdia are much larger than those of Af. inundatum and more 

dmelyln size*^°^^ which, however, they approximate 

The teown siA-hymenial layer further distinguishes this species from 
any of those so far described in these studies; it occurs also in the new 
species M stnatisporum, next to be described, which, however, has very 

charactenstic and easily recognizable conidia. 

Sub-cultures sometimes produce a wrinkled growth with very scanty 
aenm mycelium, if any at all, and usually without sporodochia. 

Myrotheciurn jollymannii is a potential parasite since, like the British 
M. roridum, It readily attacked the living leaves and stems of Viola comuta 
^amb?r°''^ suspensions were applied to detached shoots placed in a moist 

(ii) Description 

Myrothecium jollymannii sp.nov, 

tunicatis, septatis compositum. 
orbiculata, sessiha vel brevissime stipitata, cupuliformia, 0-3- 

alS,’ ^ ^ mm. diam. primo viridia demum atra, margine 

saeniiK Ton multiseptatisi crassetunicatis composite. %tae 

Sk 5-8/4 latae, septatae. Conidiophora ex 

1 P?“^=‘dibusque composita. Hyphae fertiles oHvaceo- 
brunneae, irregulanter ramosae, dense intertextae, discum compactum 

Ohservatiom on the Genus Myrothecium. N. C. Preston 273 

obscuratum efiicientes, septatae, infra septa paullo dilatatae, cellulis 
terminalibus quam basalibus brevioribus. Fhialides subhyalinae, rectae vel 
curvulae, leviter clavatae, 12-15 X2ja. Cmidia continua, pallide olivaceo- 
brunnea, cylindrica vel paullo angustata, apicibus obtusis, praecipue 
biguttulata, 8-12 X2'5/A. 

Text-fig. I. Setae. A, M. graminetm Lib.; B, M.jollymamii n.sp.; 
C, M, striatisporum n.sp. (A and B x 320; C x 1200.) 

Text-fig. 2. A, A/. gramineumlAh.: (i) spores ex Deighton M1743, (ii) spores ex Libert no. 380, 
1837; B, Af. n.sp., spores ex potato-dextrose agar; C, M, striatisporum 

spores ex cherry agar/ ^ ^ ^ ^ 



274 Transactions British Mycological Society 

Hob. Ex foliis siccantibus Nicotianae tabaci, in Nyasaland, Africa in anno 
1936 isolatum (F. W. Jollyman No. 145) et in agaro excultum et’descrin- 
turn P^s exsiccata m Herb. I. M. I. deposita sub. no. 1495 est typus 
Mycelium. Hyphae thin-waUed, colourless, 1-3/4 wide; septate cells 
usually 1 0-15/A in length. On artificial media sometimes forming a 
wnnkled growth closely appressed to the medium and consisting of tinted 

Text-fig. 3. M, jollymannii n,sp. Phialides 
from potato-dextrose agar culture. 

Text-fig. 4. M. striatispQTum n.sp. Phialides. 

Text-fig. 5 - approx, x ao. Sporodochium showing contorted 

fnngmg hyphae. Freehand drawing under Leitz ‘UltrapakV 

h5yhae about 3 /a wide of a pale brown, orange or pinkish colour, sur- 
mounted by a sparse aena,! cottony growth, always pure white, consisting 
ofslendercolomless hyphae 1-2/A in width. Sporodochia. Circuit sessile or 

Setf occasionally 2 mm. in 

, ^^een at first becoming velvety black, 
^ ^ nm of thick-walled, multiseptate, hyaline setae which at 

&st curve inwards, ^then straighten. Setae. About 100-260/A long x 5-8/A 
a blimt round^ point; usually of seven to.ten cells which 
vary from 18-26/A in length with walls about i /a thick. ConUiopfwres com- 

Observations m the Genus Myrothecium. N. C, Prestm 275 

posed of fertile hyphae and phialides (Text-fig. 3). Fertile hyphae, Olive 
brown, irregularly branched, septate, cells 8-15 x 2-4/x broadening slightly 
below the septa, the terminal cells shorter than the basal ones. Very 
closely intertwined forming a dark-coloured disk. Phialides. Sub-hyaline, 
straight or slightly curved, slightly clavate, 12-15 x2/x. Conidia. Con- 
tinuous, pale greenish brown, straight-sided, cylindrical or more usually 
very sHghtly tapering with blunt ends, the broader end flat the narrower 
end often somewhat rounded, generally biguttulate, 8-12 x 2'5ja, average 


Type isolation. From dried tobacco leaves, in Nyasaland, Africa, 1936 
(F. W. Jollyman No. 145). 

Myrotheciiim striatisporum n.sp. 

(i) Material studied and characteristic features 

This fungus was originally isolated by J. C. Neil (No. 582) from a clay 
soil in New Zealand. A sub-culture of NeiFs isolate ^as obtained through 
the Imperial Mycological Institute and this, together with subsequent 
transfers, has formed the basis of the present description. 

The sporodochia of this species (Text-fig. 5) are superficially indis- 
tinguishable from those of M. roridum^ the white Singe being composed 
of twisted hyphae among which the presence of setae is very exceptional. 
Only once have any setae been found among the many hundreds of sporo- 
dochia examined (Text-fig. I C). 

Microscopically this species is readily distinguished by its striate, or 
fluted, conidia which are generally sub-fusoid though occasionally approxi- 
mating to rod-shaped. It is further distinguishable from either M. roridum 
or M. verrucaria by the dark colour of the hymenial layer, and by the 
characteristic dark brown sterile filaments which are frequently found 
interspersed among the fertile hyphae, often extending beyond the general 
surface of the disk. 

(ii) Description 

Myrothecium striatisporum sp.nov. 

Mycelium ex hyphis hyalinis, tenuiter tunicatis, septatis 1-2 ft diam. 
compositum. Sporodochia minuta, tenuiter cupuliformia, orbicularia vel 
irregularia, o*05-0'2 mm. diam. saepe confluentia, primo viridia demum 
aterrima, margine albido ex hyphis contortis, hyalinis, tenuiter tunicatis, 
aliquanto latioribus composite. rarissimae. Conidiophora ex hyphis 
fertilibus phialidibusque composita. Hyphae fertiles brunneo-tinctae, 
praecipue 3-septatae5 irregulariter infi'a septa ramosae, ramis sursum dense 
intertextis, discum obscurum efiBcientibus; adsunt quoque hyphae steriles, 
non ramosae, atrobrunneae, inter hyphae fertiles crescentes. Phialides 
e subhyalino olivaceae, vel terminales vel e geniculo cellulae primogenitae 
oriundes, tenuissime clavatae, 7-22 x 3ft. Phialides terminales semper 
ceteris breviores. Conidia subhyalina deinde olivaceo-brunnea, continua, 
subfusoidea, apice acuta, basi minute apiculata, episporio primo laevi 
demum longitudinaliter stiiato omata, 7 x 2*5 ad 12 x 3*5^^ (av. 9*6 x gft). 

Hab. Ex terra argillacea sub agro pascuo, in Nova Zealandia in 1937 


276 Transactions British My cological Society 

isolatum (J, G. Neil 582) et in agaro excultum et descriptum. Pars 
exsiccata in Herb. I. M. L deposita sub. no. 1526 est typus. 

Mycelium, Pure white. Hyphae hyaline, slender 1-2 ja wide, thin-walled, 
septate; cells 15-25/^ long. Sporodochia, Minute shallow cups, circular or 
irregular, o*05-0'2 mm. in diameter, about 0*05-0-1 mm. deep and often 
confluent into larger masses. Composed of loosely arranged hyphae, 
arising from a compact or pseudoparenchymatous base, and terminating 
in a dense disk-like layer. Green at first, becoming jet black surrounded by 
a white rim of rather broad, thin-walled, contorted, hyaline hyphae. 
Conidiopkores composed of fertile hyphae and phialides (Text-fig. 4). Fertile 
hyphae. Tinted brown, consisting of a main axis usually of three cells with 
branches arising irregularly from below the septa and very closely inter- 
twined. Cells 3-i3ft long by about wide, the terminal cell always 
shorter than the others. Interspersed by unbranched, sterile, dark brown, 
thick-walled, somewhat sinuous filaments, 3-4/^ wide and 40ju, or more in 
length (the walls of which frequently appear verrucose). Phialides. Sub- 
hyaline to pale olive green, greenish brown in the mass, arising terminally 
or from a geniculation of the parent cell, very slenderly clavate, rather thick- 
walled, 7-22 X The terminal phialides short, straight or slightly 
flexuose, the lateral ones longer and bent sharply near the base, the whole 
thus forming a closely packed, even, hymenial layer. Conidia. Small, 
continuous, sub-fusoid, guttulate, broadest slightly below the middle, with 
the apex pointed and the base terminated by a minute stalk-like pro- 
tuberance less than ifx long. Epispore at first smooth becoming fluted, 
usually with about fifteen ridges arranged longitudinally and often some- 
what spirally. At first sub-hyaline, when mature smoky olive brown, 
7 X 2*5-i2 X 3*5jn, average 9*6 x 3 jit. 

Hah. Clay soil. New Zealand. 

The author wishes to thank the Director of the Imperial Mycological 
Institute for access to material in the herbarium and especially Mr E. W. 
Mason for his unfailing interest and helpful criticism; he is also greatly 
indebted to Miss E. M. Wakefield for the Latin diagnoses of the new 


Libert, M.-A. (1837). P/. Crypt. Ard. Fuse, rv, No. 380. 

Preston, N. C. (1943). Observations on the Genus Myrotkecium Tode. I. The three 
classic species. Trezwj, Brit. myc. Soc. xxvi, 158-68. 

Plate XV 

Fig. I. Lib. ex Herb. R. B. G. Kew. Spores, x 1000. 

Fig. 2. M. gfamirmtmiUb. (Deighton M. 1743) ex Herb. 1 . M. L Spores, x rooo. 

Fig. 3, M. Jollytmnnii n.sp. Sporodochia on agar, showing fringing setae, x 32. 

Fig. 4. M. Jollyrnamdi Fragment of spor^ochium with setae, x 460. 

Plate XVI 

Fig. 5. n.sp. Gonidiophores, X 1000. 

Fig. 6 . M. striatisporum n.sp, SmEill sporodochia showing fringing hyphae, X 60 approx. 

Mg. j. M* striati$porum n^^ Spores, x 1000. 

Fig. S. M, striatisporum n.sp. Spores, showing striations, x 3000 approx. 

{Accepted for publication 2^ May 

[ 277 ] 



By G. T. INGOLDj Birkbeck College^ University of London 
(With 2 Text-figures) 

To a perithecium from which spores are being actively discharged there 
must presumably be a steady supply of water making good the loss by 
evaporation so that the turgidity of the ripening asci may be maintained. 
Probably most of the lignicolous stromatal Pyrenomycetes (e.g. Nectria 
spp., Diatrype spp., Endothia spp. and Hypoxylon spp.) depend directly on 
rain for this water supply and spore discharge is limited to periods during 
or immediately after rain when the fungal tissue is temporarily turgid. 
Daldinia concentrica (see Ingold, 1946) is exceptional in being able to dis- 
charge spores during prolonged dry periods by virtue of the considerable 
reserve of water in the stromatal tissue. 

Epichloe typhina (Fr.) Fr., the choke of grasses, causes a systemic 
infection of its host and eventually produces its perithecial stroma, during 
July and August, as a crust around the unexpanded leaves immediately 
above a node of the stem. The innermost hyphae of the stromatal tissue are 
in intimate association with the living cells of the host. The experiments 
reported in this paper were designed to answer the following question: does 
the fungus during periods of active discharge obtain its water from the host 
cells and, therefore, indirectly from the transpiration stream of the grass, or 
is discharge limited to damp periods when the stroma is directly wetted by 
rain or dew? 

The specimens used were collected at Sevenoaks, Kent. The form of the 
fungus on Dactylis glomerata was finally selected for use because in the 
material available the long thread-like ascospores showed no tendency to 
break up into part-spores and the units in the spore deposit were 190 x 2 /i.. 
On the other hand, from specimens of the fungus on Holms mollis the units 
measured 57 x 2ft no doubt due to the breaking up of the ascospores into 
shorter part-spores. The process appears to be carried still further in the 
form on Agrostis tenuis since the spore deposit from this strain was composed 
of units 32 X 2ft. Each measurement of length is the average of 1 00 units of 
the spore deposit chosen at random. It would be interesting to know if 
these differences obtain in other localities and if the forms on other grasses 
show still further variations. 

Each diseased shoot of Dactylis used for an experiment consisted of a 
length of stem, above this a stroma of Epichloe^ and above this again a single 
green leaf (Fig. i). Each shoot was severed from the plant by a cut made 
below water to avoid the interruption of the transpiration stream by air 
blocks in the vessels. With their cut ends in water the shoots were brought 
indoors for study. 

278 Transactions British Mycological Society 

It was found that from the fungus on such shoots, freely exposed to the 
dry air of a room in summer, spore discharge continued for several days so 
long as the cut ends of the shoots remained in water. 

Discharge can clearly be seen by the naked eye if, when the sun is 
shining brightly, the fungus is held more or less between the observer and 
the sun, but in such a way that the stroma can be viewed against a dark 
background. Then the individual spores, each about a fifth of a millimetre 
long, can be seen as needle-like motes in the sunbeam. They first come into 
view about half a millimetre away from the stroma and then are slowly 
carried away by gentle air currents. If discharge is very vigorous, there are 
so many spores in the air around the stroma at any instant that it is im- 
possible to observe discharge from a single ascus, but if spores are not being 
liberated in too great numbers, the successive discharge of the eight spores 
from an ascus, which is completed in much less than a second, can be seen. 
The eight discharged spores, maintaining roughly their formation like 
a squadron of mi nute aeroplanes, can often be followed for a few centi- 
metres as they drift away. 

X F G H I 

1 / /. 






! / , 

' ■ 

! ^ 1 

[ — ^ i 



Fig. I. Method of measuring rate of spore discharge from stroma of Epkhloe, A, cork support; 
specimen tube; C, glass blocks; D, wooden block; F, glass slide; F, stem of grass; G, node 
of grass; stroma of Epickloe; /, le^ of grass. 

In studying the rate of spore discharge the arrangement shown in Fig. i 
was used. The cut end of the shoot dipped into water in a specimen tube 
clamped by a rubber band in a more or less horizontal position to a special 
cork support (d) consisting of a horizontal and a vertical limb. The stem of 
the shoot passed through a hole in the vertical limb and through another 
hole in the cork of the specimen tube (B). In both holes the shoot was 
wedged firmly with dry cotton-wool. The free leaf of the shoot was held 
secmely in position between two blocks of glass (C). The stroma in this 
way occupied a horizontal position and below it a wooden block (D) was 
placed. For observing the rate of spore discharge a microscope slide {E) 
was used with a square centimetre, sub-divided into square miUimetres, 
etched upon its upper surface. This could quickly be slipped into position 
under the fungus and it was so arranged that at each observation the etched 
square occupied exactly the same position relative to the stroma. After 
a brief exposure (usually fifteen or thirty seconds) the slide was removed 
and the spores deposited on the square counted under the low power of the 
microscope. These short exposures were necessary, since it is difiicult to 
count the spores if there are too many, and a single stroma when most 
active may discharge 2000-10,000 spores a minute. The apparatus was not 
covered, but was freely exposed to the air of the room, the door and windows 

Water-^relations of Spore Discharge in Epichloe; C. T. Ingold 279 

being closed to avoid draughts. The limitations and errors of the method 
are obvious, but it can be used to give a reasonably accurate measure of 
changes in the rate of spore discharge. The effect on discharge of inter- 
rupting the transpiration stream was studied by cutting the stem with 
a razor at the point marked X (Fig. i). 

Using the method described above, certain facts were clearly demon- 
strated. In the first place, there tends to be a daily periodicity of discharge 
with the minimum in the morning and a maximum in the afternoon or 
evening. This was observed in nearly all the specimens studied. Secondly, 
considerable fluctuations in the rate of discharge occur over short intervals 

Fig. 2. A and B, curves of rate of spore discharge from two different stromata of Epichloe plotted 
against time (g.m.t.). One experiment ( 2 l) performed on 23-24 July, the other (B) on 
18-19 1 94 ^* Observations of temperature (dotted line) and relative humidity (inter- 

rupted line) are for 23-24 July and should be considered in connexion with A, Arrows, X 
and T, indicate the time when the stem of the host plant was severed, thus interrupting the 
transpiration stream. 

of time. Thirdly^ and most important, cutting the stem at the point Zwhen 
the rate of discharge is at a high level rapidly leads to a cessation of spore 
liberation. Theresults of two experiments aye represented diagraimnatically 
in Fig. 2 . Both were conducted on fairly warm July days. In both, cutting 
the stem at a time when the rate of discharge was high and apparently 
increasing led to a rapid fall of the rate to zero after an hour or so. 

These observations indicate that spore discharge can be maintained in 
fairly dry air without any external supply of water to the stroma, but that 
the fungus is dependent, like the host tissue with which it is so intimately 
associated, on the water of the transpiration stream and when this is inter- 
rupted by cutting the stem, the rate of spore discharge soon falls to zero. 

28 o 

Transactions British Mycological Society 


Spore discharge from Epichloe typhina can be observed with the unaided eye. 
A method is described for studying the rate of spore liberation. 
Discharge tends to be periodic with a minimum in the morning and a 
maximum in the late afternoon or evening. 

In fairly dry air spore discharge continues for several days provided 
that the water supply to the host tissue is maintained, but when the 
transpiration stream is stopped by severing the stem, discharge soon ceases. 


Ingold, G. T. (1946). Spore discharge in Daldinia concentrica, Trans. Brit myc, Soc. xxdc, 
43 - 51 - 

{Accepted for publication 3 May 1947) 



Microbiologist, Mushroom Research Association, Taxl^, Hunts 
(With i Text-figure) 

The ohief problem which confronts the student of soil fungi is the discrimi- 
nation between fungi which are present in the soil as normal active 
inhabitants and those which occur there as accidental inactive inhabitants. 
While the agents responsible for their introduction and distribution in the 
soil are to some extent the same, for example, percolating water and 
organisms of various kinds (earthworms, eelworms, insects, etc.) their 
ultimate behaviour is different in so far as the former grow and reproduce 
in the soil, while the latter die or remain dormant in the state in which 
they were originally introduced as spores or pieces of mycelium. 

Waksman (1932), whose immense amount of work on soil organisms is 
well known to workers in this field, advocates two methods of obtaining 
fungi from soil — the direct and indirect methods. The direct method which 
consists in incubating for 24 hr. particles of soil on a nutritive agar, is claimed 
by him to have the advantage that growth from already active mycelium is 
obtainable within this period. He points out, however, that on account of 
the different rates at which various fungi grow, only the faster-growing 
species may be collected in this way. His indirect method is selective only 
in so far as various media are used in the plating out of the soil dilutions. 
It does not eliminate the accidental contingent of the soil fungus 

Blair (1945) briefly reviewed the various techniques used for soil fungus 
studies and their applications up to 1945. The majority of those which he 
mentioned were either some modification of the soil-suspension method or 
of the contact-slide method. He himself used for growth studies of Rkizoc- 
tonia mycelium in the soil, what he termed a modification of the Rossi- 
Cholodny slide technique. Before immersing the microscope slides in soil 
he inoculated them with portions of cultures of Rhizoctoniay zud subse- 
quently estimated the growth of mycelium over the slide after fixation and 
staining. While the slide method is adaptable for the purpose of estimating 
mycelium quantitatively it is, however, only of limited value for qualitative 

As far as I am aware, Chesters (1940) was the first to devise an apparatus 
(immersion tube) which contained culture medium and wHch could be 
immersed in the soil for the purpose of obtaining fungi actively gromng 
there. This method enabled him to isolate and grow the fungi so obtained. 
Unfortunately, at least for the average worker, the apparatus is difficult to 

282 Transactions British Mycological Society 

Recently^ in studies on fungi growing in the casing soil of mushroom 
beds and in the underlying compost, as well as in soil incubated in beakers, 
I have obtained satisfactory results with a very simple device. It consists 
of two standard hanging-drop slides which are clipped together so that 
their concavities lie opposite each other. Enclosed between the concavities 
is a small quantity of suitable agar medium (Fig. la^ b). A culture 
chamber is thus provided to which fungi can gain access by growing in 
between the slides from the surrounding soil. The moisture of condensation 
which collects in drops between the slides while the latter are immersed 
in the soil enables growth to take place until the agar is reached. 

Among the fungi which so far have been collected by this method are 
species of Alternaria^ Botryotrichum, Cylindrocarpon^ Fusarium and Trichoderma, 

Although the potentialities and adaptability of this device have not yet 
been fully explored, the author is sufficiently persuaded of its usefulness to 
recommend it to those engaged in work on soil fungi. 


c d e 

Fig. I. 

Preparation and use of slide-traps 

The laboratory bench is swabbed with alcohol where the slide-traps are to 
be prepared. 

Hanging-drop slides are cleaned and matched in pairs so that their 
concavities He in apposition. They are separated and laid on the bench 
in* two parallel rows proximal and distal to the operator, and a sterile Petri 
dish Hd is laid base upwards on the bench between the individuals of 
each pair. 

The slides in the proximal row are, successively, dipped in alcohol, 
flamed, and placed concavity upwards, one under each Petri dish Ud. 

A small quantity of sterile melted agar medium, e.g. soil extract agar 
such as that used by Ellis (1940), is quickly pipetted into the concavity of 

SlideHtaps for Soil Fungi. C.J. La Touche 283 

each of the slides under the Petri dish lids. When 'the agar has solidified, 
the counterparts from the other row of slides are sterilized as already 
described, and each is then inverted over its fellow so that the agar is 
enclosed between the two concavities. 

The slides may be secured together by means of various types of clip, as 
illustrated (Fig. ir, dy e)y or simply with antiseptic plasticine. 

When collections are made in mushroom beds, the surface soil for a 
depth of \ in, is scraped away with a sterile knife. The slide-traps are flamed 
quickly and pushed edgewise into the soil vertically or horizontally until 
completely buried. The surface soil is then replaced. When collections are 
made in casing soil in beakers, a layer of several inches of soil is first put 
into the beakers, the traps are partly immersed in this, and are then 
covered over completely with more soil: this is done with every possible 
precaution to avoid contamination. 

After a suitable period of immersion, usually a week in soil incubated in 
beakers at 22° C., and a fortnight to 3 weeks in the mushroom beds (where 
lower temperatures prevail) the traps are lifted and examined under the 
microscope. Hyphae will usually have grown in between the slides across 
their width, and may be traced from the edge of the slides right into the 
agar. If the agar has not been reached by the hyphae the traps may be 
incubated in a moist chamber until growth is well established in the agar. 
When the agar has become invaded by mycelium, the slides are prised 
open and the agar is removed quickly with the sterile flattened tip of 
a nichrome wire and placed on sterile agar medium such as 3 % malt 
extract agar in a Petri dish. When the agar is invaded from two opposite 
sides it may first be cut in half with a micro-scalpel and the two halves 
then removed separately. The removal of the agar from the traps is carried 
out preferably in a sterile glass-fronted chamber which admits only the 
arms for the purpose of manipulation. Isolations may be made subse- 
quently at leisure from the Petri dish transfer. 

The advantages of the above method in my opinion are as follows : 

(1) It requires a minimum of apparatus and technique. 

(2) The amount of agar medium used in the slide-traps is infinitesimal. 

(3) A large number of slide- traps may be set up quickly for use in a 
small area or volume of soil. 

(4) The vegetative and often even reproductive structures, may be 
studied in situ without disturbance. 

(5) Contamination by bacteria is usually avoided. 

(6) If the slides composing the trap are kept tightly apposed, marauding 
organisms, capable of carrying spores and bacteria, are kept out. 

(.7) Isolates may be made easily by transferring the whole or part of the 
agar to media in Petri dishes. 

The disadvantages are: 

(1) If several different kinds of mould grow into the same small volume 
of agar some may be suppressed and lost owing to cbmpetition. 

(2) Flooding of the surrounding soil may result in water penetrating 
between the slides by capillary attraction and contaminating the agar with 
bacteria or mould spores held in suspension. 

284 Transactions British Mycological Society 

The author wishes to acknowledge with thanks the permission of tlie 
Director of Research and the Directors of the Mushroom Research 
Association to publish this article. 


Blair, I. D. (1945). Techniques for soil fungus studies. N.Z* Sci, Tech, xscvi A, 258-71. 
Chesters, G. G. G. (1940). A method of isolating soil fungi. Trans, Brit, myc. See, xxiv, 


Ellis, M. (1940). Some Fungi isolated from Pinewood soil. Trans, Brit, myc. Sac, xxiv, 

Waksman, S. E. (1932). Principles of Soil Microbiology, 

{Accepted for publication 28 April 1947) 

[ 285 ] 


By ROBERT yiolLKYy Department of Plant Pathologp^ 

University College^ Dublin 

In the autumn of 1946 Apple Scab was severe in the orchard at Glasnevin^ 
Dublin^ and the fruit on four large trees of the variety Annie Elizabeth was 
so badly diseased that it was never picked. This fruit remained hanging on 
the trees throughout winter until the last week in January 1947, when with 
the onset of severe wintry weather it was attacked by birds and every 
apple was knocked down into the grass underneath. Here, a heavy fall of 
snow protected the fruit from being completely devoured. The snow 
remained throughout February and part of March, and after its disap- 
pearance it was found that while many of the fallen apples had been 
reduced to mere skins, by slugs and birds, others still retained their shape 
more or less. Examination of the old skins and particularly of the shrunken 
fruit on 25 March 1947 showed a fairly prolific development of perithecia 
of the scab fungus Venfuria inaequalis. The majority of the asci and asco- 
spores were only in the process of formation at this time, but some were 
fully mature. Perithecia on the fruit are not so obvious as those occurring 
on dead leaves, but they can be readily identified with the aid of a pocket 
lens or binocular microscope in the laboratory. 

The literature on Apple Scab is voluminous, and numerous observers in 
both hemispheres have found perithecia of Venturia inaequalis on dead over- 
wintered leaves, but apparently there is no previous record of perithecial 
development on scabbed fruit. 

{Accepted for publication 28 April 1947) 

[ 286 ] 


By T. fetch 

A list of the entomogenous fungi of Great Britain was published in Tram, 
Brit rnyc. Sac. xvii (1932), 170-8. Since then a number of additions have 
been made, largely through the investigations of Mr E. A. Ellis, on the 
Broads in the neighbourhood of Norwich, which have yielded several 
species, new or new to Britain. The value of the new records has been 
enhanced by many identifications of the host insects, some of which have 
been kindly furnished to Mr Ellis by Captain N. D. Riley, of the British 
Museum (Natural History). To them, and to the numerous correspondents 
who have sent me specimens, I tender my most sincere thanks. 

The localities given in the previous list are not repeated, except where 
only one or two occurrences are known. Some apology is needed for the 
inclusion of so many records, but they serve to indicate the range and 
frequency of the species and any variation in hosts. Where a species is said 
to be common, only records of special interest are included. Names 
marked with an asterisk are additions to the previous list. 


EmpiLsa Muscae Cohn, in Nova Acta Acad. Caes. Leopold.-CaroL Germanicae 
Nat. cur. XXV (1855), Abh. i, 301. 

On flies. Generally distributed, and common out-of-doors in the summer, 
usually on grasses. Frequent in gardens on Asparagus beds. On a Capsid 
{Lygus pabidinus)^ Cambridge, 19 August 1933 (F. T. Brooks). 

Empusa Culicis A. Braun, Algarum unicell. gen. nov. (1855), p. 105. 

On gnats. Hornsea, 16 July 1931. On small fly. Holt House Wood, 
King’s Lynn, 17 August 1931. 

Empusa Aulicae in litt. (cf. Bail in Schrift d. naturf. Gesellsck. zu 
Danzig^ n.f., ii (1869), 3). 

On caterpillars. On grass, North Wootton, 16 October 1933. On 
lettuce, North Wootton, 22 September 1939. On Brassica, North Wootton, 
16 October 1941. 

Empusa Tipulae Fetch, in Trans. Brit. myc. Soc. (1932), 170; 

Entomophthora Tipulae Fres., in Abhandl. d. Senchnberg. naturf. Gesellsck., ii 
(1858), 206. 

On a Tipulid, Aldborough, E. Yorks., 12 September 1933; resting spores 
in the abdomen, spherical, smooth, hyaline or pale yellow, 32-45 ft 
diameter, wall up to 4^ thick. Studland, Dorset, October 1936 (E. W. 

A Revised List of British Entomogenous Fungi. T. Fetch 287 

Empusa Fresenii Nowak., in Proc. Krakow Acad. ScL (1883), p. 153. 

On aphid {Macrosiphum taraxaci Kalt.) on dandelion, Hapton, Lancs., 
6 September 1925 (H. Britten). 

Empusa Planchoniana (Cornu) Fetch, in Trans. BriL n^c. Soc. xxi (1937), 36; 
Entomophthora Planchoniana Cornu, in Bull. Soc. Bot, France^ xx (1873), 189; 
Entomophthora ferruginea Phillips, in Ann. Mag. Nat. Hist.^ Ser. 5, xviii 
(1886), 4; non Empusa Planchoniana Thaxt., Entomophthoreae US. (1888), 
p. 165. 

On aphids. On greengage. North Wootton, 30 May 1935. Deepdale, 
Barnard Castle, September 1933. On Pentatrichopus fragariae Theob. on 
strawberry, Gian Conway, 7 June 1943 (Dr C. A. Wood). 

Empusa Thaxteriana Fetch, in Trans. Brit myc. Soc. xxi (1937), 36; Empusa 
Planchoniana Thaxt., Entomophthoreae U.S. (1888), p. 165. 

The. entries in the previous list under Empusa Planchoniana (Cornu) Thaxt. 
should be referred to this species. 

On aphids on Prunus Padus, Deepdale, Barnard Castle, September 1933. 
On Homopteron [Philaenus leucophthalmus)^ Upton Broad, 20 July 1942, 
Wheatfen Broad, 13 August 1942 (E. A. Ellis). 

Empusa papillata Thaxt., Entomophthoreae US. (1888), p. 166. 

On flies {Sciara sp.) on Lactarius^ Common, E. Yorks., 

2 August 1936. 

^Empusa apiculata Thaxt., Entomophthoreae US. (1888), p. 163. 

On flies and a leaf-hopper, Heatree, Dartmoor, 24 September 1935. 

"^Empusa Tenthredinis (Fres.) Thaxt., Entomophthoreae US. (1888), p. 162; 
Entomophthora Tenthredinis Fres., in Abhandl. d. Senckenberg. naturf. Gesellsch. 
II (1858), 205. 

On larvae of sawflies on Spiraea Ulmaria, Parish Marsh, Wheatfen Broad, 
14 and 27 June 1943 (E. A. Ellis). 

* Empusa Grylli (Fres.) Nowak., in Proc. Krakow Acad. Sci. (1883), p. 168; 
Entomophthora GrylliPies^'m. Bot. Z'^tung,'siv 

On a grasshopper, Allerthorpe Common, E. Yorks., 2 August 1936. 

* Empusa Acaridis Fetch, in Trans. Brit. myc. Soc. xxvn (1944), 87. 

On a mite {Pergamasus crassipes Linn.), Home Marsh, Wheatfen Broad, 
9 August 1942 (E. A. Ellis). 

*Empusa Forficulae (Giard) Fetch var. major Fetch, in Trans. Brit. myc. Soc, 
xxvn (1944), 87. 

On earwigs. Old Lakenham, 10 August 1942 (E. A. Ellis). 

288 Transactions British Mycological Society 

Entomophthora sphaerosperma Fres., in Bot. Z^itung^ xiv (1856), 882. 
Common. Usually on aphids and flies. 

On Ichneumonidae, Deepdale, Barnard Castle, September 1933; Beck- 
dale near Helmsley, 3 September 1935 ; Millington, E. Yorks., August 1936, 
September 1937. On an ant, Duncombe Park, Helmsley, 2 September 
^935* On Hemiptera on nettles, Haye Park, Ludlow, September 1937. 
On nymph of Heteropteron, Upton Broad, 20 July 1942 Ellis). On 

nymph of Homopteron, Parish Marsh, Wheatfen Broad, 14 June 1943 
(E. A. Ellis). Epidemic on larvae of Pieris brassicae^ Adisham, Kent, 
5 September 1945 (S. G. Jary). 

The record in the previous list, on beetle, Hornsea, 14 July 1931, was 

Entomophthora Aphidis Hoffm., in Fres. in AbhandL d. Senckenberg. naturf. 
Gesellsch. ii (1858), 208. 

On aphids. Common. On root aphis of lettuce {Pemphigus lactucarius)^ 
September 1937 (N. C. Preston), with resting spores (see Trans. Brit, myc, 
Soc, XXIII (1939)5 128). 

^Entomophthora occidentalis Thaxt., Entomophtkoreae U.S. (1888), p. 171. 

On aphids. On sycamore, Millington Wood, near Pocklington, Yorks., 
4 September 1937. On Ranunculus repens^ Old Lakenham Marshes, 26 July 
1942 (E. A. EUis). 

Entomophthora Aphrophorae Rostrup, in Botan. Tidskr. xx (1896), 128; 
Fetch in Trans. Brit. myc. Soc. xix (1935)5 179. 

On Homoptera, usually Philaenus spumarius {Aphrophora spumaria L.). 
Arncliffe Woods, near Whitby, August 1930. Grassington, September 
1931. Aldborough, E. Yorks., September 1933 and August 1937. Hedon, 
E. Yorks., September 1933 and August 1937. Lartington and Deepdale, 
N.W. Yorks., September 1933. Dipton Wood near Corbridge, September 
1933. Beckdale near Helmsley, September 1935. Pallathorpe, Bolton 
Percy, July 1937 (W. G. Bramley). Millington Wood near Pocklington, 
September 1937. The Sneap, Co. Durham, October 1937 (A. W. Bartlett). 
Fritton Bog, Suffolk, September 1933 (E. A. Ellis). On Athysanus grisescens 
Zett., Aldeifen Broad, Norfolk, July 1935 (E. A. Ellis). Poor's Marsh, 
Wheatfen Broad, September 1942 (E. A. Ellis). On Philaenus leucopktkalmuSy 
Tas Marshes, Thurston, and Upton Broad, July 1942 (E. A. Ellis). Wych- 
wood Forest near Oxford, August 1940 (E. W. Jones). 

Entomophthora muscivora Schroet., in Krypt. ’-Flora v. Scklesien, m (1886), pt. i, 

On flies. On ivy, Aldborough, E. Yorks., 30 September 1932. Balders- 
dale, N.W. Yorks., 18 September 1933. Allerthorpe, 3 August 1936. 
Hubberholme, 8 September 1936. Little Bud worth near Tarpoley, 
Cheshire, ii September 1938 (F. B. Stubbs). Newport, Salop, 15 October 
1945 (N. G. Preston). Wytham, Berks., 9 October 1946 (Prof. G. D. H. 

A Revised List of British Entomogenous Fungi. T. Fetch 289 

Entomophthora americana Thaxt., Entomophthoreae U.S. (1888), p. 179. 

On flies. Eavestone Woods, Ripon, August 1933 (Miss L. M. Anderson). 
Baldersdale, N.W. Yorks., 18 September 1933. Deepdale, N.W. Yorks., 
19 September 1933. Newcastle, October 1935 (A. W. Bartlett). Doncaster, 
September 1936 (Mrs Morehouse). Hubberholme, 8 September 1936. 
Park Wood, Elland, 15 July 1937 (E. Bearing). Plymouth, September 
1942 (per Dr G. B. Williams). Godstow Osier Holt near Oxford, June 
1945 (E. W. Jones). 

The record of this species on a leaf-hopper in the previous list is an error. 

A • 

Entomophthora variabilis Thaxt., Entomophthoreae US. (1888), p. 183. 

On flies. Lartington, 16 September 1933, ^'esting spores pale brown, 
spherical, smooth, 30-40 /x- diameter. Beckdale near Helmsley, 3 September 
1935. Allerthorpe, 2 August 1936. Buckden, September 1936. 

Entomophthora dipterigena Thaxt., Entomophthoreae U.S. (1888), p. 177, 

On flies. Common. Helmside Ghyll, Dent, 5 June 1933 (F. A. Mason), 
resting spores external, hyaline, spherical, thick-walled, smooth, 26-33^ 
diameter. Lartington, 16 September 1933. Dovedale, Derbyshire, i June 
1935. Buncombe Park, Helmsley, i September 1935, resting spores 
2 7-40 ft diameter. Heatree, Dartmoor, 24 September 1935. Allerthorpe 
Common and Millington Wood, August 1936. Buckden, September 1936. 
North Wootton, 24 September 1936. On Calliphora vomitoria, Norwich, 
13 September 1936 (E. A. Ellis). Haye Park, Ludlow, 21 September 1937. 
Boyne Park, Ludlow, 23 September 1937. On Psychodidae^ Wheatfen 
Broad, 7 July 1941 and 27 June 1943 (E. A. Ellis). 

Entomophthora echinospora Thaxt., Entomophthoreae U.S. (1888), p. i8o. 

On flies. Lartington, 16 September 1933. Buckden, 5 September 1936. 
Millington Wood, 4 September 1937. Buttercrambe Moor Wood, 6 Sep- 
tember 1937- Boyne Park, 23 September 1937. Dartington, 25 September 
1935. Burntfen Broad, 25 September 1941 (E. A. Ellis). Wheatfen Broad, 
13 August 1942 (E. A. Ellis). North Wootton, 22 August 1942, mass of 
resting spores bright coral red, not yellow. 

[Entomophthora Lampyridarum Thaxt., Entomophthoreae U.S. (1888), p. 169. 
The record of this species in the previous list is an error.] 

Entomophthora coleopferorum Fetch, in Trans. Brit. myc. Soc. xvii (1932), 172, 
and xxvii (1944), 88. 

On beetles. Lartington, Yorks., 16 September 1933. On Sitonesjlavescens, 
Wick, 10 October 1937 (Miss D. J. Jackson). 

^Entomophthora anglua Fetch, in Trans. Brit. myc. Soc. xxvii (1944), 89. 

On beetles. On Hornsea, 14 July 1931. On Agriotes sputator, 

Coton, Cambs., 3 May 1931 and June 1933 (E. W. Jones). On Cantharis, 
Pickering, 6 June 1938 (W. G. Bramley). On Lochmaea suturalis^ Edinburgh, 



290 Transactions British Mycological Society « 

June 1939 (R. W. G. Dennis). On Cantharis sp.. Old Lakenham, 25 June 

1942 (E. A. Ellis). On a Staphylimd, Wheatfen Broad^ 9 August 1942 
(E, A. Ellis). On larvae of Galerucella tenella^ Wheatfen Broad, 14 June 

1943 (E. A. Ellis). Roydon Common, Norfolk, May 1945 (C. P. Fetch). 

"^Entomophthora virescens Thaxt., Entomophthoreae US. (1888), p, 178. 

On a caterpillar on Brassica, North Wootton, 30 September 1942. 

Entomophthora {Tarichium) Richteri (Bres. & Star.) Bubak, in Ann. Myc. xiv 
(1916), 341 ; Massospora Richteri Bres. & Staritz, in Hedwigia (1892), p. 133; 
Tarichium Richteri (Bres. & Star.) Lakon, in Z^itscL /. PjianzenKr. xxv 
(1915), 257; Entomophthora Lauxaniae Bubak, in Hedwigia (1903), p. (100.) 

In flies. Recorded in the previous list as Entomophthora Lauxaniae. 
Rokeby, 17 September 1933. Baldersdale, 18 September 1933. Deepdale, 
N.W. Yorks., 19 September 1933. Austwick, Yorks., 18 September 1934. 
Buckden, September 1936. 

Entomophthora [Tarichium) atrosperma Fetch, in Trans. Brit. myc. Soc. xvii 
(1932), 172. 

On aphis. Grassington, 12 September 1931 (F. A. Mason). 

* Tarichium megaspermum Cohn, in Beitr. z- Biol. d. Pflanzen i (1875), 58. 

In larvae of Plusia gamma on Pisum, Romney Marsh, Kent, July 1946 


*Myrophagus ucrainicus (Wize) Sparrow, in Mycologia, xxxi (1939), 443; 
Olpidiopsis ucranicaV^vzt, in Akad. Umiejet. Krakow [Bull. Intern. Cl. Sci. Math, 
nat.) (1904), 715; Entomophthora reticulata Fetch, in Trans. Brit. myc. Soc. 
XXIII (1939), 127; Jfaturalist (March 1940), p. 68. 

In a dipterous pupa, Ingleborough, September 1934. 


*Torrubiella aranicida Boud., in Rev. Myc. vii (1885), 227. 

On large spiders under hanging moss on a cliff face, Hubberholme, 
Yorks., 8 September 1936; on spider, Wheatfen Broad, 2 August 1945 
(E. A. Ellis). 

*Torrubiella albolanata Fetch, in Trans. Brit. myc. Soc. xxvn (1944), 85. 

On spiders among Cladium, etc., usually Gongylidium rufipes, Wheatfen 
Broad, 21 June 1942, 9 and 13 August 1942, 14 June 1943 (E. A. Ellis). 
On spider’s egg cocoon, Wheatfen Broad, 15 April 1945 (E. A. Ellis). 

Conidial stage — Cylindrophora aranearum Fetch, in Trans. Brit. myc. Soc. 
xxvn (1944), 85, with the perithecial stage or separately. 

*Torrubiella albotomentosa Fetch, in Trans. Brit. myc. Soc. xxvn (1944), 86. 
On a dipterous pupa in a dead stem of Glyceria maxima, Parish Marsh, 

A Revised List of British Entomogemus Fungi. T. Fetch 291 

Wheatfen Broad, 20 May 1943 (E. A. Ellis). On dipterous pupae. Wheat- 
fen Broad, 17 May 1945 (E. A. Ellis). 

Cordyceps militaris (L.) Link, Handb. in (1833), 347. 

On lepidopterous larvae and pupae. Generally distributed. On larvae 
of Hepialus^ Austwick, 18 September 1934. 

The conidial stage of this species is not Spicaria {Isaria) farimsa^ but a 
Cephalosporium (see Fetch, Cordyceps militaris and Isaria farinosa, in Trans. Brit, 
myc. Soc. xx (1936), 216-24). This has been named Cephalosporium militare 
by Kobayasi in The Genus Cordyceps and its Allies (1941)5 p. 113. 

Cordyceps gracilis Mont. & Dur., Flor. Alger, i (1846-9), 449. 

On larvae of Lepidoptera, usually Hepialus. Ludlow, 28 May 1932. 
WhitclifFe Woods, Richmond, Yorks., 24 April 1937 (J. B. Nicholson). 
Ponterwyd near Aberystwyth, 2 May 1940 (S. J. Hughes). Long Crendon, 
Oxon., March 1945 (J. Howard Brown), 

Conidial stage — Isaria dubia Delacroix. 

Cordyceps entomorrhiza (Dicks.) Link, Handb. ni (1833), 347. 

On a beetle larva, Bulstrode, Bucks., in the autumn (Dickson, ante 1 785) . 
No further records. The records for N.E. and N.W. Yorks, in Mason and 
Grainger, Catalogue of Torkshire Fungi^ p. 33, were not in the MSS. sub- 
mitted to me, and doubtless refer to Cordyceps gracilis. 

Conidial stage — probably Hirsutella eleutheratorum (Nees) Fetch, but until 
the perithecial stage is available, that cannot be ascertained. 

"^Cordyceps tuberculata (Lebert) Maire, in Bull. Soc. d^HisL nat. d. VAfrique du 
JSford^ viii (1917)5 165; Acrophyton tuberculatum Lebert, in ^eitschr. f. wiss. 
Zool. IX (1858), 448; Torrubia Sphingum (Schw.) TuL, SeL Fung. Carp, m 
(1865), 12; Cordyceps Sphingum (Schw.) B. & C., Fungi Cubenses, no. 746 

On larvae, pupae and imagos of Lepidoptera. On a pupa in moss under 
nettles, Alderfen Broad, Norfolk, 15 August 1940 (E. A. Ellis). On a pupa 
under alders, Low Common, Hellington near Norwich, 9 May 1943, with 
the conidial stage, Akanthomyces Sphingum (Schw.) Fetch, on the same pupa 
(E. A. Ellis), On lepidopterous pupae, Alderfen Broad, 19 August 1945, 
Strumpshaw Marsh, 10 October 1943, Brundall Marshes, 16 August 
1945 (all coll. E. A. Ellis). 

Previous records of Cordyceps Sphingum in Britain have been based on 
Berkeley and Broome’s record of the conidial stage, as Isaria Sphingum Schw., 
on pupae of Diptera, Kincardineshire in Notices of British Fungi^ no. 1710. 
That specimen, however, is a linear form of Spicaria {Isaria) farinosa. 

Cordyceps sphecocephala (Kl.) Godk^^ Vegetable WaspSy ttc. {i^^ 2 )yp. 40. 

On Hymenoptera. The Wood, Wheatfen Broad, 20 September 1942, 
found unattached among dead leaves (E. A. Ellis). 


292 Transactions British Mycological Society 

Cordyceps Forquigmni Qjuelet, in Assoc. Franc, p. PAvancement d. Sciences (1887), 

p. 6, 

On flies. Aucliincmive, Ayrshire, 1934 (R. W. G. Dennis). On an 
Anthomyid, Cyrtoneura estabulans Fin., The Wood, Wheatfen Broad, 
26 March 1942 (E. A. Ellis). Spreacombe, Braunton, North Devon, 
September 19455 July 1946 (F. R. Elliston Wright). 

Conidial stdigc—Hymemstilbe muscaria Fetch. 

Cordyceps clavulata (Schw.) E. & E., JSforth American Pyreno (1892), p. 61; 
Cordyceps pistillariaeformis B. & Br., in Ann. Mag. Mat. Hist. Ser. 3, vii (1861), 

45 ^* 

On scale insects, Lecanium spp. On a scale insect on wych elm, Batheaston, 
Somerset, March i860, October i860 (Broome). No further records. 
Conidial stage — Hirsutella lecaniicola (Jaap) Fetch. 


Myriangium Duriaei Mont. & Berk., in HookePs London Journ. Bot. iv (1845), 

73 - 

In Britain, usually on Chionaspis salicis on ash in the south and west. 
No further records. 


^Oospora ovorum Trabut, in Rev. gen. Bot. iii (1891), 404. 

On cocoons of a Braconid parasite on Odonestis potatoria, Wheatfen 
Broad, 23 September 1945 (E. A, Ellis). 

Scopulariopsis penicillioides (Delacr.) A. L. Smith & Ramsbottom, in Trans. 
Brit. myc. Soc. v (1915), 164; Monilia penicillioides Delacr., in Bull Soc. myc. 
France^ xiii (1897), 114. 

On pupae of the Cinnabar Moth from Farnham Royal, May 1931. 

^Acremonium tenuipes Fetch, in Trans. Brit. myc. Soc. xxi (1937), 66; Sporo-- 
trichum aranearum Cav., Fung. Longob. exsicc. no. 240 (1892); Verticillium sp., 
Sartory, Sartory & Meyer, in C.R. Soc. Biol. Paris, cvii (1931), 53“5* 

On spiders and spider egg masses. Reffley Wood near King’s Lynn, 
8 June 1930. Holt House Wood near King’s Lynn, 17 August 1931. The 
Wood, Wheatfen Broad, 7 March 1943 (E. A. Ellis). 

"^Acremonium aranearum Fetch, in Trans. Brit. myc. Soc. xvi, 242. 

On Homopteron, Fostwick Marshes, 16 August 1945 (E. A. Ellis). 

^Cylindrophora aranearum Fetch, in Trans. Brit. myc. Soc. xxvri (1944), 85. 

The conidial stage of Torrubiella albolanata Fetch. On spiders, with the 
perithecial stage or separately. Wheatfen Broad, 9 August 1942, etc. 
(E. A. Ellis). 

A Revised List of British Entomogemus Fungi, T, Fetch 293 

^Cephalosporium [Acrostalagmus) aphidicola Petch^ in Trans, Brit, myc, Soc, xvi 
(1931), 71. 

On aphid {Capitophorusfragariae) on strawberry, Auchincruive, Scotland, 
February 1939 (C. E. Foister). 

Cephalosporium {Acrostalagmm) LefroyiTLovnc^ in Gard, Chron, LVii (1915)5 139. 
On Aleyrodes vaporarium^ c£ Fetch in Trans, Brit, myc, Soc, xxv (1942), 262. 

'^Cephalosporium {Acrostalagmus) subclavatum Fetch, in Trans, Brit, myc, Soc, 
xxv (1942), 262. 

On lepidopterous larvae. On a Caradrinid larva from a fen locality, 
May 1941 (J. C. F. Fryer). On larvae oiNudaria senex Hb,, Home Marsh, 
Wheatfen Broad, 9 August 1942 and 19 May 1943 (E. A. Ellis). 

Cephalosporium {Acrostalagmus) coccorum Fetch, in Trans, Brit, myc, Soc, x 

(1925). 175- 

On Chionaspis^ Lepidosaphes^ etc. On subterranean coccids, Hagg Wood, 
Burnsall, 15 September 1931. On mealy bugs, Stratton Strawless, 
4 October 1934. On mealy bug on gorse, Totnes, 27 September 1935. 

Cephalosporium {Acrostalagmus) Eriophytis (Massee) Fetch, in Trans, Brit, myc, 
Soc, XVI (1931), 66; Botrytis Eriophyes Massee, in Taylor, Journ, econ, Biol, 
IV (1909), 5; Verticillium Eriophytis (Massee) Sacc. & Trott., Syll, Fung, xxn 
(1913), 1299. 

On Eriophyes ribis in ‘big bud’ of black-currant. 

Cephalosporium {Acrostalagmus) muscarium Fetch, in Naturalist (April 1931), 
p. 102. 

On flies. Deepdale, N.W. Yorks., 19 September 1933; Marsh Cote, 
Saltaire, 24 October 1933 (W. P. Winter). On Urophora solstitialis L., 
Cambridge, March 1935 (F. T. Brooks). Dartington, 25 September 1935. 
Cambridge, 31 July 1942 (F. T. Brooks). 

On aphids. Bolton Woods, 17 September 1931; Lartington, 16 Sept- 
ember 1933; Rokeby, 17 September 1933; Deepdale, 19 September 

On beetles. Deepdale, 19 September 1933; Pallathorpe, 26 August 1935 
(W. G. Bramley); on Plateumaris braccata, Wheatfen Broad, 9 August 1942 
(E. A. Ellis); on a weevil {Erirrhinus nereis Pk.), Wheatfen Broad, 4 June 
1942 (E. A. Ellis). 

On Homoptera. Wheatfen Broad, 25 July 1935 (E. A. Ellis); Old 
Lakenham Marshes, 26 July 1942 (E. A. Ellis). 

On Ichneumon. Wheatfen Broad, 9 August 1942 (E. A. Ellis). On 
chalcid larva {Torymus cyanimus), Cambridge, March 1935 (F. T. 
Brooks) . 

On spider cocoon. Hubberholme, 9 September 1936. On spider, 
Wheatfen Broad, 9 August 1942 (E. A, Ellis). 

294 Transactions British Mycological Society 

Cephalosporium {Acrostalagmus) dipterigenum Fetch, in Naturalist (April 1931), 
p. 102. 

On a beetle {Cantharis lateralis), Studland, June 1933 (E. W. Jones). On 
a spider, Millington Wood near Pocklington, 4 September 1937. On 
Homoptera, Wheatfen Broad, 5 October 1944 and Old Lakenham Hall, 
27 August 1942 (E. A. Ellis). 

* Cephalosporium militare Kobayasi, The Genus Cor dy ceps and its Allies (1941), 
p. 113. 

The conidial stage of Cordyceps militaris (L.) Link. At the base of the 
perithecial clavae of the Cordyceps, Austwick, Yorks., 18 September 1934. 
Also obtained in culture from the ascospores of the Cordyceps, cf. Fetch, 
Cordyceps militaris and Isaria farinosa, in Trans. Brit. myc. Soc. xx (1936), 
216-24. Kobayasi did not give a description in the publication cited. 

Beauveria Bassiana (Bals.) Vuill., in Bull. Soc. Bot. Fj'ance, lix (1912), 40. 

On insects of all kinds, among moss or dead leaves, or under bark or 
logs, etc. Numerous records, of which the following are noteworthy. On 
a bee {Bombus pratorum), Castle Gardens, Norwich, 2 October 1933 (E. A. 
Ellis). On spiders. North Wootton, 18 October 1933. On Ichneumon, 
North Wootton, 18 October 1933. On fly. Pinhole Cave, Derbyshire, 
24 March 1934 (F. A. Mason). On Picromerus bidens, Camberley, May 1934 
(E. E. Green) . On Hepialus larvae, Harpenden, April 1 934 (H. F. Barnes) . 
On leaf-hoppers, Rawcliffe near Goole, 4 August 1934. On a wasp. North 
Wootton, I March 1935. On beetles, Pallathorpe, Bolton Percy, 26 August 
1936 (W. G. Bramley). On Homopteron, Upper Hellesdon, 25 August 
1940 (E. A. Ellis). 

Beauveria densa (Link) Picard, in Ann. Serv. Epiph. i (1913), 160. 

On insects of all kinds, but usually on beetles. On a weevil, Rawcliffe 
near Goole, 4 August 1934. On beetles {Strophosomus lateralis, etc.), Belton 
Heath, Suffolk, 3 January 1935 (E. A. Ellis). [On cockchafer larvae and 
adults, Kennington forest nursery near Oxford and Fairoak nursery near 
Tintern, 1 942 (J. M. B. Brown) see note.] On a caterpillar, Old Lakenham 
Hall, 2 August 1942 (E. A. Ellis). On a beetle {Plateumaris braccata), 
Wheatfen Broad, 9 August 1942 (E. A. Ellis). On beetles {Galerucella 
tenella), Wheatfen Broad, 25 February 1943 (E. A. Ellis). On a wasp, 
Wheatfen Broad, 25 February 1943 (E. A. Ellis). On ants, Hellington, 
10 June 1943 (E. A. Ellis). 

* Beauveria effusa (Beauv.) Vuill., in Bull. Soc. Bot. France, lix (1912), 40. 

On pupa of Agrotis ripae (larva from Studland), June 1936 (C. Diver). 
On indet. insect, Wheatfen Broad, 20 December 1942 (E. A. Ellis). On 
beetles {Galerucella tenella), Wheatfen Broad, 6 May 1943 (E. A. Ellis). 

Metarrhizium Anisopliae (Metsch.) Sorok,, in Z^itschr. d. Kais. Landw. 
Gesellsch,f.Neurussland{i^']^),p. 9 .b&. 

On insects of all kinds. Isolated from a dead wireworm, Cambridge, 

A Revised List of British Entomogenous FungL T. Fetch 295 

October 1939 (R. Leach). On wireworm, Rothamsted, January 1941 
(H. C. Gough), On larva of wireworm {Corymbites cupreus), Cardiff, 
I September 1942 (S. J, Hughes). 

^Cylindfodendmm suffultum Fetch, in Traris Brit, myc. Soc, xxvii (1944), 91. 

On pupae of Diptera. On pupae of Clytocerus ocellaris Meig., Trowse 
Marshes, 7 August 1942 ; Wheatfen Broad, 9 August 1942, 13 August 1942, 
19 June 1942. On Tipulid pupae, Wheatfen Broad, 19 June 1943. On 
Stratiomyid pupae, Wheatfen Broad, 19 June 1943. On dipterous pupae, 
St Olave's, Suffolk, 18 July 1944 (all coll. E. A. Ellis). 

^Veriicillium menisporoides Fetch, n.sp. On spider, St Olave^s, Suffolk, 
18 July 1944 (E. A. Ellis). 

"^Spicaria prasina (MaubL) Saw., in Descr. Cat Formosan Fungi (1919); 
Momuraea prasina MaubL, in Bull, Soc, myc, France^ xix (1903), 295. 

On caterpillars. On a caterpillar on grass, Wheatfen Broad, August 
1939; Bradeston Hills, 30 September 1945; Cringleford Marshes, 28 October 
1945 (all coll. E. A. Ellis). 

"^Spicaria stricta Fetch, in Trans, Brit, rnyc, Soc. xxvn (1944), 89. 

On a spider, Wheatfen Broad, 13 August 1942 (E. A, Ellis). 

^Spicaria fumoso-rosea (Wize) VassiL, in Morbi Plant,, Leningrad, xviii 
(1929), 1 13; Isariafumoso-rosea 'Wizt, in Bull, Acad, Cracovie (1905), p. 354; 
Fetch, in Trans, Brit, myc, Soc, xxvn (1944), 90. 

On larvae of Lampyris noctiluca (glow-worm) in marsh litter. Wheatfen 
Broad, 12 September 1937, 13 August 1942, 25 August 1942, 27 September 
1942, 15 October 1944; Blundeston Marshes, Suffolk, 4 September 1943; 
Strumpshaw Marsh, Norfolk, 9 October 1944, 12 August 1945; Alderfen 
Broad,* 19 August 1945 (E. A. Ellis). On a lepidopterous pupa, Wheatfen 
Broad, 4 October 1942 (E. A. Ellis). 

Spicaria (Isaria) farinosa (Holms.) VuilL, Bull. Soc, myc, France, xxvn (1910), 
75“~82; Isaria farinosa (Holms.) Fr., Syst. Myc. iii (1832), 271; Spicaria 
{Isaria) Swantonii (A. L. Smith) P^tch., m. Naturalist (April 1931)5 p. 103; 
Coremium Swantonii A. L. Smith, in Trans, Brit, myc, Soc, xi (1919), 156. 

On insects of all kinds, either as an Isaria or a simple Spicaria, 

Aspergillus depauperatus Fetch, in Trans. Brit, myc. Soc. xvi (1931), 244. 

On Lepidosaphes ulmi on hawthorn, Hunstanton, February 1921. 


Isaria dubia Dclacr., in Bull, Soc, myc, France, ix (1893), 264. 

The Gonidial stage of Cordyceps gracilis. Grown in culture from ascospores 
oi Cordyceps gracilis, May 1932. On larvae of Swift Moth {Hepialus) from 
Dr Redington, Glos., g November 1940. 

296 Transactions British Mycological Society 

*Isaria ochracea Boud., in Bull. Soc, myc. France, xix (1903), 197. 

On Sphinx ligustri, Norwich, 1939. 

*Isaria tenuipes Peck, Thirty-first Report N.T. State Museum, p. 44 (1879). 
On lepidopterous pupae. The Wood, Wheatfen Broad, 15 October 1938, 

18 August 1940, 18 September 1941, 20 September 1942 (E. A. Ellis). 

Hymenostilbe muscaria Fetch, in Naturalist (April 1931), p. loi. 

The conidial stage of Cordyceps Forquignoni. On flies. Hubberholme, 
9 September 1936. Whiteclifie and Boyne Park, Ludlow, September 1937. 
On a Cordylurid [Norellia upinimana Fin.). Wheatfen Broad, 26 March 1942 
(E. A. Ellis). On Anthomyids, Wheatfen Broad, 18 June 1942, i October 
1942 (E. A. Ellis). Spreacombe, N. Devon, December 1944, 30 August 
1945, September 1945 (F. R. Elliston Wright). 

On unweathered specimens the conidia measure 7-1 1 x 3-4-5 /r. 

Hymenostilbe arachnophila (Ditm.) Fetch, in Naturalist (August 1931), p. 249; 
Isaria arachnophila Ditm., in Sturm, Deutschl. Flora, Abt. m (1817), Bd. i, 
III, PI. 55. 

On spiders, usually attached to living leaves. Hunstanton, July 1932 
(T. Harris). Lartington, 16 September 1933. Deepdale, N.W. Yorks., 

19 Septemljer 1933. Dipton Wood near Corbridge, 21 September 1933. 
Chopwell Wood, Durham, 22 September 1933. Whitecliffe and Kinlet 
near Ludlow/September 1937. Old Lakenham, 25 June 1942 (E. A. Ellis). 
Bradeston Hills, 4 October 1945 (E. A. Ellis). 

Prof. E. B. Mains has found a Cordyceps, C. Thaxteri Mains, in company 
with H. arachnophila in America {Journ. Elisha Mitchell Sc. Soc. isv (1939), 1 20. 

* Hjmenostolbe sphecophila (Ditm.) Fetch, in Trans. Brit. myc. Soc. xxi (1937), 
55; Isaria sphecophila Ditm., in Sturm, Deutschl. Flora, Abt. in (1817), 
Bd- L i55> Bl- 57- 

On Hymenoptera. The conidial stage of Cordyceps sphecocephala. On 
Ichneumon, Chopwell Wood, Durham, 22 September 1933. On Ich- 
neumon, Horsey Mere, 5 May 1938 (E. A. Ellis). 

In the previous list, mention was made of a probable specimen of 
Hymenostilbe sphecophila figured by Worthington G. Smith in Card. Chron. 
on a bee, later said to be a wasp. In a more detailed figure in W. G. Smith, 
Diseases of Field and Garden Crops (1884), p. 59, the insect is clearly a wasp. 

Akanthomyces Sphingum (Schw.) Fetch, in Trans. Brit. myc. Soc. xxvii (1944), 
82. Isaria Sphingum Schw., Syn. Fung. Carol. Super. (1822), p. 126; Akantho- 
myces aculeata Lebert, in ^eitschr.f. wiss. ^ool. ix (1858), 448; Hymenostilbe 
(Schw.) Fetch, in Trans. Brit. myc. Soc. xvi (1932), 2 i‘]', l Isaria 
floccosaFr., Syst. Myc. in (1832), 274; ? Isaria strigosa Fr., Syst. Myc. ni (1832), 

On lepidopterous pupae, larvae and images. The conidial stage of 
Cordyceps tuberculata. On a lepidopterous pupa, with the Cordyceps, Low 
Common, Helhngton near Norwich, 9 May 1943 (E. A. Ellis). 

A Revised List of British Entomogenous Fungi. T, Fetch 297 

Gibellula aranearum.iSchw,) Syd., Fetch in Naturalist (August 1931)5 p. 249 ; 
Isaria aranearum Schw., Sjn. Fung. Carol. Super. (1822)5 p. 126; Gibellula 
aranearum Syd^j in Englefs BoL Jahrb. lvii (1922), 321, 

On spiders attached to grasses or to the underside of leaves. Bradwell, 
SufFolk5 18 July 1933. Fritton Bog, Lothingland5 3 September 1933 
(E. A. Ellis). Baldersdalej 18 September 1933. Aldeby, E. Norfolk^ 7 July 
1935 (E. A. Ellis). Wheatfen Broadj 25 July 1935, 5 October 1941, 
22 October 1944 (E. A. Ellis). Berry Pomeroy, 26 September 1935. 
Hubberholme, 8 September 1936. Burntfen Broad, 25 September 1941 
(E. A. Ellis). Hellington, 10 June 1943 (E. A. Ellis). On Gong^lidium 
mfipes^ Wheatfen Broad, 27 June 1943 (E. A. Ellis). Brundall Marshes, 
24 August 1945 Ellis). On a Homopteron and a small dipterous 

pupa, Swannington Upgate Common, 30 July 1944 (E. A. Ellis). 

Hirsutella Eleutheratorum (Nees) Fetch, in Naturalist (February 1932), p. 46; 
Isaria Eleutheratorum Nees, System d. Pilze (1817), p. 85. 

On beetles and beetle larvae. Dunscombe Park, Helmsley, i September 
1935. Buckden, 7 September 1936. On Rhizophagus ferrugineus Pk., 
Ballater, 18 August 1938, and Glendye, Scotland, 20 August 1938 (H. S. 
Hanson). On Paederis riparius, Wheatfen Broad, 5 October 1941, Hickling 
Broad, 10 July 1945 (E. A. Ellis). On Staphylinids, Surlingham Broad, 
12 July 1944, Postwick Marshes, 18 October 1945, 28 October 1945 (E. A. 
Ellis). Strumpshaw Broad, 31 August 1944 (E. A. Ellis). Cirencester, 
February 1944 (— Shaw, per Miss E. M. Wakefield). 

^Hirsutella Saussurei (Cooke) Speare, in Mycologia^ xii, 69. 

On Ichneumonid and small Hymenoptera, Postwick Marshes, 6 May 
1945, iSOctober 1945, 28 October 1945 (E. A. Ellis). On ?, Alderfen Broad, 
19 August 1945 (E. A. Ellis). 

^Hirsutella citriformis Speare, in Mycologia^ xii, 70. 

On Homoptera (including Kelisia scottii (Fieb.), Areopus pulckellus Curt.) 
and Heteroptera {Nabis lineatus Dahlb.), Postwick Marshes, 18 October 
1945 (E- A. Ellis). 

Hirsutella subulata Fetch, in Naturalist (February 1932), p. 48. 

On caterpillar, Milton, Northants. (J. Henderson, without date, but 
before 1881), recorded as Isaria fioccosa Fr. in B. & Br., Notices of British 
Fungi, no. 1096. No further British records, but has been found in America, 
on the larva of the ‘ Codling Moth ’ under a piece of bark, Washington, 
D.C., and on the larva of Aegeriapyri Harris in apple bark, Flatbush, New 
York; cf. Fetch in Trans. Brit. myc. Soc. xxi (1937), 57. 

*Hirsutella exoleta (Fr.) Fetch, in Naturalist (November 1936), p. 251; Isaria 
exoleta Vv., Syst. Myc. m (1832), 275; Cordycepsfuliginosa Ces., Comment. Soc. 
Critt. Ital. i {i 86 i), Sj. 

On lepidopterous pupae. Steeton Whin near Tadcaster, July 193® 
(W. G. Bramley). Hubberholme, September 1936. 

298 Transactions British Mycological Society 

Hirsutella acridiorum Fetch, in Trans. Brit. myc. Soc. xvn (1932), 177. 

On a cricket, Lartington, 16 September 1933. On a Homopteron, 
Aldeby, E. Norfolk, 7 July 1935; Surlingham Broad, 15 July 1944; Post- 
wick Marshes, 16 August 1945 (E. A. Ellis). 

* Hirsutella Aphidis Fetch, in Naturalist (March 1936), p. 60. 

On aphids. Aldborough, E. Yorks., 12 September 1933. Brundall, 
27 September 1943 (E. A. Ellis). 

* Hirsutella dipterigena Fetch, in Trans. Brit. myc. Soc. xxi (1937), 53. 

On flies [Blepharoptera serrata) in caves. Pinhole Cave, Cresswell, Derby- 
shire, 24 March 1924 (Leslie Armstrong). In a ‘stalactite cave’, Yeal- 
hampton, Devon, June 1906 (Herb. B.M.). 

*Hirsutella lecaniicola (Jaap) Fetch, in Trans. Brit. myc. Soc. xvra (1933), 53. 

The conidial stage of Ophiacordyceps clavulata (Schw.) Fetch. With the 
perithecial stage on Lecanium on wych elm, Batheaston, Somerset, March 
and October i860 (Broome). 

Syngliocladium aranearum Fetch, in Trans. Brit. myc. Soc. xvn (1932), 177. 
On a spider among dead leaves, St Leonard’s Forest, Horsham, 31 May 


* Syngliocladium Cleoni (Wize) Fetch, in Trans. Brit. myc. Soc. xxv (1942), 
265; Acremonium Cleoni Wize, in Bull. Acad. Sci. Cracovie (1905), p. 352. 

Onwireworms (Agriotes sp.), Rothamsted, February 1941 (H. C. Gough), 
with the sclerotial stage, Sorosporella uvella (Krass.) Giard. On larvae of 
wireworms [Corymbites cupreusB.), Cardiff, 26 August 1942 (S. J. Hughes). 


*Aegerita insectorum Fetch, in Trans. Brit. myc. Soc. xxi (1937), 63. 

On larva of Urophora solstitialis L. (Diptera), Cambridge, March 1935 
(F. T. Brooks). On larva oiStenichneumon trilineatus, St Andrews, iSJanuary 
1936 (Miss D. J. Jackson). 

Microcera coccophila Desm., in Ann. Sci. nat. Ser. 3, x (1849), 359. 

In England, usually on Chionaspis salicis on willow in the south-west. No 
further records. 


[Cladosporium Aphidis Thuem., in Oester. Bot. Z^itschr. xxvii (1877), 12. 

This is Cladosporium herbarum Link, and, on aphids and flies at least, it 
usually follows an attack oi Empusa or Entomophthora\ cf. Fetch in Trans. 
Brit. nyc. Soc. XIX igo.] 

A Revised List of British Entomogenom Fungi. T. Fetch 299 

Fungi parasitic on entomogenous fungi 

Melanospora parasitica TuL, Selecta Fung, Carp, iii, 10. 

On Spicaria (Isaria) farinosa^ RawclifFe near Goole, 4 August 1934. On 
Beauveria Bassiana^ Dartingtonj 25 September 1935. On ? on beetles 
{Plateumaris braccata), Wheatfen Broad, 9 and 13 August 1942 (E. A. Ellis). 
On Isaria tenuipes^ Wheatfen Broad 20 September 1942 (E. A. Ellis). On 
Beauveria densa on a caddis fly, Wheatfen Broad, 18 October 1942 (E. A. 

^Sphaeroderma fusisporum Fetch, in Naturalist (March 1936), p. 58. 

On Spicaria {Isaria) farinosa^ Hirst Wood, Saltaire, September 1935 
(W. P. Winter); North Wootton, 18 October 1936. On Syngliocladium 
Cleoni on wireworms from Rothamsted, February 1941. 

^Stilbella ramosa (Peck) Fetch, in Trans, Brit, myc, Soc, xxi (1937), 53; 
Stilbum ramosum Peck, Bull, Soc, Nat. Sci. Buffalo^ 1 (1873), 69; Stilbella 
Kervillei (Quelet) Lingelsh., in Ber, Deutsch, Bot. Gesell. xxxix (1921), 149; 
Stilbum Kervillei Quelet, in Gadeau de Kerville, Bull, Soc. Amis d. Sc, Nat, 
Rouen (1884), pi. i (extr.); Stilbella Arndtii Lingelsh., in Ber, Deutsch, Bot, 
Gesell, XXXIX (1921), 149; Stilbella setiformis (Vahl) Fetch, in Trans, Brit, 
myc, Soc, xvm (1933), 55; Clavaria setiformis Vahl, in Naturhist, Selsk. Skrivt. 
n (i793)> 50 - 

Parasitic on species of Hirsutella and on Cordyceps which have a Hirsutella 
conidial stage. On Hirsutella dipterigena on Blepharoptera serrata^ Cresswell 
Caves, Derbyshire, 1923 (F. A. Mason in Journ. Bot, (August 1931), 
p. 205) ; Pinhole Gave, Cresswell, 24 March 1934 (Leslie Armstrong) . 

It is probable that the transfer to Stilbella has been made in America 
prior to that cited above. 

Sporotrichum Isariae Fetch, in Naturalist (April 1931), p. 102. 

On Spicaria farinosa on gnats. Pinhole Cave, Cresswell, April 1934 (Leslie 
Armstrong). On Spicaria {Isaria) farinosa^ RawclifFe near Goole, 4 August 
1934. On Spicaria {Isaria) farinosa^ Upton Broad, 8 August 1940 (E. A. 
Ellis). On Spicaria {Isaria) farinosa on Homopteron {Acocephalus nervosus)^ 
Old Lakenham, Norwich, 18 October 1938 (E. A. Ellis). On Aphrophora 
alniy Wheatfen Broad, 20 April 1939 (E. A. Ellis). 

New species 

Vertidlliimi menisporoides Fetch, n.sp. 

On a dead spider, submaritime fen, St Olave’s, Suffolk, 18 July 1944 
■:(E. A. Ems).. , 

Spider covered by a rather loose greyish weft, from which arise erect 
conidiophores, especially along the legs. Conidiophores up to 6oo/x high, 
gregarious but distinct; stem hyaline, 3-4 /x diameter below, 2/x diameter 

^00 Transactions British Mycologtcal Society 

above, septate, rigid, bearing whorls of phialides and, towards the base, 
short lateral branches, long, with an apical group of phialides. 
Phialides four or five in a whorl, conoid, tapering to the apex, not septate, 
25-30 ju, long, 2*5 p diameter below. Conidia apical, hyaline, continuous, 
persisting parallel in ovoid clusters, cymbiform or arcuate, produced at 
each end into a long point, 12-20 x 2-2 -5 with some narrow-oval, ends 
not produced, 9-12 x 2p, in the same cluster. 

Mycelio laxo cinereo araneam obducente; conidiophoris ad 600/^ alt., 
gregariis sed non fasciculatis, stipite hyalino, septato, rigido, infra 3-4 /x 
diam., supra 2(1 diam., phialides verticillatos, et ad basin ramulos breves 
ferentibus; phialidibus quatuor v. quinque verticillatis, conoideis, ad 
apicem attenuatis, 2 5-30 /x long., infra 2-5/1 diam.; conidiis apicalibus, 
hyalinis, in fasciculus parallele persistentibus, cymbiformibus vel arcuatis, 
utrinque longe attenuatis, 12-20 x 2-2*5/c, vel anguste ovalibus, 9-12 x 2ja, 
non attenuatis. 

On a spider, Suffolk, England. 

Notes on the foregoing records 
Entomophthora sphaerosperma Fres. 

An epidemic of Entomophthora sphaerosperyna on caterpillars oi Pieris 
brassicae on Brassica in Kent ^ was reported to the Plant Pathology 
Laboratory, Harpenden by Mr S. G. Jary in September 1945. The details 
in the following paragraph are taken from Mr Jary’s report. 

The caterpillars were observed to be dying in one corner of a field of 
15 acres on 31 August and by 5 September the disease had spread over the 
whole field. The caterpillars turned brownish, some showed signs of 
mycelium, and others were entirely covered by mycelium and fastened by 
it to the leaves. Many failed to pupate, and numbers of pupae were 
obviously dead, and disintegrated when touched, being filled with a 
brownish fluid. Near the corner of the field, where the disease was first 
observed, it seemed that almost every caterpillar had been killed, very few 
caterpillars or pupae being found on trees. 

Larvae of Pieris brassicae are the standard (original) host of Entomophthora 
sphaerosperma, and epidemics of the disease have been reported on several 
occasions on the Continent, but they have not been previously recorded in 
this country. On the other hand, E, sphaerosperma, as at present understood, 
is one of our commonest species Entomophthora and occurs on insects of all 
kinds, though I had never before seen it on caterpillars. There is, however, 
a record of its occurrence on a small green caterpillar on grass ( Trans. BriL 
myc. Soc. vi, 202), and in September 1946 I received specimens on the 
larvae of Plutella maculipennis from Scotland per Dr G. E, Foister. They 
were of normal consistency, and bore the typical primary and secondary 
conidia, but no resting spores. On the other hand, the specimens received 
from Mr Jary were squashy, with a fringe ofhyphae fastening them to the 
leaf and an external yellowish crust of resting spores, but primary and 
secondary conidia and cystidia could not be found, the whole mass being 
more or less disorganized. 

A Revised List of British Entomogemus Fungi. T. Fetch 301 

The original description of Entomophthora sphaerosperma by Fresenius was 
apparently based on specimens in a similar condition to those from Mr Jary 
— hence the inappropriate epithet. The disease was subsequently in- 
vestigated by Brefeld, who also worked with Pieris larvae and apparently 
added details of the primary and secondary conidia. But the rarity of 
epidemics on Pieris larvae in this country, in spite of the abundance of the 
fungus on other hosts, and the difference in its effect on these other hosts, 
suggest that there is more than one fungus included under this name. It 
should be ascertained whether the primary conidia and cystidia occur on 
Pieris larvae in an early stage of the disease, before the production of the 
resting spores. 

Tarichium megaspermum Cohn. 

This species was described by Cohn in 1875. It occurred in Germany in 
the larvae of Agrotis segetum which were turned black by the spores, and the 
disease has been styled the Black Muscardine. It does not appear to have 
been observed again in Europe before its present occurrence, but it has 
been recorded on cutworms at the Manitoba Agricultural College, Canada. 
It has not been redescribed, and is known only from Cohn’s description and 
figures. Cohn’s description, as summarized in Thaxter’s monograph, stated 
that ' The resting spores are apparently azygospores borne laterally or 
terminally from hyphae and are peculiar on account of their dark brown 
epispore which is marked by sinuous furrows. The epispore is also fre- 
quently opaque showing no furrows. The spores are spherical and of large 
size measuring from 34 to 55 (jl, average 50/x.’ Other available translations, 
e.g. Winter in Rabenhorst, Cooke, Gueguen, make no mention of smooth 

What appears to be this species was forwarded to me from the Plant 
Pathology Laboratory, Harpenden, having been collected by Mr S. G. 
Jary in larvae of Plusia gamma on Pisum^ Romney Marsh, Kent, at the end 
of July 1946. The spores are spherical or slightly ellipsoid, black or black- 
brown, 36~50 /x diameter. They are at first loosely contained, singly or in 
pairs, in a pale brown, thin-walled, smooth mother cell, which is ellipsoid 
or produced at one end into a cylindrical or conico-cylindrical appendage. 
They soon become free, and numerous empty mother cells occur among 
the black spores. The actual spore is black, but the external black material 
appears to be a deposit, not a spore wall. Beneath the black substance is 
a thin, brown, smooth epispore, followed by a hyaline inner wall, 3/x thick. 
The mother cells are evidently produced terminally or laterally on hyphae 
as figured by Cohn, but in no instance have I seen this cell wall furrowed, 
and the wall of the spore beneath the black deposit is smooth. I would 
suggest as a possible explanation that the spore wall observed by Cohn 
was the wall of the mother cell, artificially wrinkled by drying or some 
other means. 

It has been supposed that Tarichium megaspermum k the azygospore stage 
of an Entomophthora, and the name Tarichium has been used generically for 
resting spores (real and alleged) of Entomophthoraceae. But the prpent 
specimens suggest that T. megaspermum should be placed in the Chytridiales. 

302 Transactions British Mycological Society 

I have received, per the Imperial Mycological Institute, specimens on 
lepidopterous larvae, Kenya, collected by Dr R. M. Nattrass, Nairobi, 
December 1945, matching those from Romney. 

Oospora ovorum Trabut. 

In this specimen the fungus formed minute, scattered, white, loose, 
pulvinate tufts, or clavate, isarioid clavae up to 0-25 mm. high, o*i2 mm, 
diameter above, 0*05 mm. diameter below, with globose or slightly oval 
conidia, 0*8-1 /x diameter, in long branching chains, terminal on hyphae 
i/x diameter. Compare Tram, Brit. myc. Soc. xxm, 143. 

Ammonium aranearum Fetch. 

Mycelium in this specimen loose, floccose ; hyphae 3 fi diameter, collapsing 
in sections ; phialides lateral on the hyphae, scattered or clustered, base oval, 
3‘5“"4M high, 2 [I diameter, with a short sterigma, collapsing into a hair-like 
thread; conidia globose, 3/x diameter, or oval, 3-4 x 2 *5-3 ft, rather thick- 
walled. This has the same structure as Acremonium album Preuss. It would 
seem that a new genus might be established for these species of Acremonium 
which have collapsing phialides, 

Beauveria densa (Link) Picard. 

In 1 935-^6, I received the Le Moult culture of Isaria densa from two 
sources and was surprised that the fungus should have been placed in 
Beauveria^ but after examination of old cultures of other species of Beauveria 
I concluded that it had 'run out’. In 1942, however, I received specimens 
on cockchafer adults and larvae from Mr J. M. B. Brown which agreed 
with the Le Moult culture. It appears therefore that the latter has not 
'run out’, and that it is not a Beauveria. The fungus (or fungi) on cock- 
chafers requires further investigation, which I regret I cannot now under- 

Isaria fumoso-rosea (Wize) Vassil. 

In recording this species for Britain in Trans. Brit. myc. Soc. xxvii, go, 
I stated that the spores of the British specimens were white in mass, and 
therefore the record was doubtful, as the fungus is known in Russia as the 
Pink Muscardine. Mr E, A. Ellis has since sent me further specimens on 
glow-worms, all of which are decidedly pink, except in one collection in 
which they vary from faintly pink to white. Presumably the pink colour 
fades. - 

Isaria ochracea '&ou 6 .. 

This specimen, on an imago o^ Sphinx ligustri^ was found dead in Norwich 
and taken to the Museum. The under surface bears numerous pale brown, 
short, laterally flattened clavae, with some taller and expanding above into 
a brush, but not forming a head. The conidia now present are oval, 
3-4 X I •5-2 /x, in chains. Clusters of prophialides and phialides occur 
along the hyphae of the clava, the prophialides being globose, 3 jtx diameter, 
and the phialides narrow flask-shaped, 5-6 x 2 /x. The conidiophore has the 
same type of structure as in Isaria tenuipes dixid I. ochracea . {vls figured by 

A Revised List of British Entomogenous Fungi. T. Fetch 303 

Boudier). The conidia differ in shape from those of L tenuipes, but are the 
same shape as those of L ochracea, Boudier, however, gave the size of the 
conidia of the latter as 6-8 x 4-4*5/i. 

Hirsutella subulata Fetch* 

This species was described in 1932 from a specimen in Herb. Kew. 
collected before 1881, and the only conidia found measured 2'5xi/x* 
Later two American specimens in the Farlow Herbarium, one dating from 
1890, were found to have conidia 4-6 x and 4 x respectively 

{Trans. Brit mjc. Soc. xxi, 123). More recently, Miss V. K. Charles, 
working with fresh material in America, has found the conidia to be 

Hirsutella Saussurei (Cooke) Speare. 

This fungus, which occurs frequently on wasps and hornets in tropical 
countries, was found on several occasions by Mr E. A. Ellis on Ichneu- 
monidae and small Hymenoptera on marshes near Norwich during May- 
October, 1945. The clavae were usually short, compared with tropical 
examples, but on one specimen they were up to 15 mm. long. In four 
collections, the conidia were cymbiform or narrow-oval, 6-9 x 2-2 *5 /x. 

Speare, who examined specimens on Polistes in Hawaii, and others on 
Polistes spp. from North Carolina, California and the West Indies, stated 
that this species was readily distinguished by its long, narrow and usually 
allantoid spores, and he gave their dimensions as 9-1 1 x There 

would seem a possibility, therefore, that ^Hirsutella Saussurei\ as at present 
understood, is a ‘collective species’. The following spore measurements 
were made on some of the specimens I have examined during the last 
sixteen years. I give the dates of collection and examination, as they may 
have some bearing on the results obtained. 

On Polistes^ Fiji, allantoid or cymbiform, 6-7 x i*5/x; 1923; Nov. 1930. 
On wasp, Panama, narrow-oval or cymbiform, 3-5 x Feb. 1917; 

Nov. 1930. 

On small Hymenopteron, Trinidad, narrow-oval, 3*5-5 x Oct. 

1916; Nov. 1930. 

On Hymenopteron, Trinidad, cymbiform, 5-6 x i*5/x; Jan. 1919; Nov. 

On wasp, Trinidad, oblong-oval or allantoid, 4-7 x i-i-5jLx; Jan. 1921; 
Feb. 1931. 

On Hymenopteron, Venezuela, cymbiform, 6-7xi*5-2/x; Aug. 1932; 
April 1934. 

These specimens, in order, were Williams, nos. 42, 43, 44, 66; Wake- 
field, no. 44; and Venezuelan Fungi, no. 1443. 

The conidiophore of a Hirsutella is very fragile, and unless treated care- 
fully the head of conidia is readily broken off. Perhaps the larger heads 
are lost first. But it would seem possible that the conidia may increase in 
size after abstriction, while in the globule of mucus which forms the head. 
However, the conidia in the British specimens of H. Saussurei are more 
broadly cymbiform than in the tropical forms. 

304 Transactions British Mycological Society 

Hirsutella citriformis Speare. 

A number of examples of this species, which is usually associated with 
warmer countries, were found by Mr E. A. Ellis on a marsh near Norwich 
on HomopterUy including Kelisia scotti (Fieb.) and Araeopus pulchellus Curt., 
and a Heteropteron, Nabis lineatus Dahlb, The somewhat rigid, hair-like, 
brown clavae were up to 20 mm. long. 

As far as I am aware, this species has not been previously reported in 
Europe. A fungus of similar appearance, Isaria stilbiformis, was described 
by Spegazzini from a specimen on a small Pentatomid in Italy. Spegazzini 
described his species as 3-7 mm. high, the hyphae of the compound stem 
diverging at the apex and forming a subglobose or pyriform head, the apices 
of the hyphae being clavate, and each bearing a cylindrical, subacute, 
hyaline conidium, 8-1 1 x 2-2 -5 /-t. There seems little doubt that the fungus 
was a Hymenostilbe. Typically, the basidia of a Hymenostilbe are situated 
uniformly along the clava, but I have seen examples in which they were 
collected into a terminal head in H, melanopoda and H, formicarum. 

Mr Ellis points out that these specimens of Hirsutella citriformis^ collected 
October 1945, came from the same locality as his specimens of H, acri- 
diorum^ collected August 1945, and suggests that these may be forms of the 
same species. In H, eleutheratorum, a mucedinous form was found in the 
catacombs of Paris and was named Isaria Guignardi Maheu, but there does 
not appear to have been any similar difference in environment of the 
collections recorded here, which both occuri'ed in living leaves. 

(Accepted for publication 28 January 1947) 


^Cordyceps memorabilis Ces., Comm, Soc, Critt. ItaL i, (1861), 16. 

On a carabid larva, Wheatfen, Norfolk, 10 September 1947 (E. A. Ellis). 

[ 305 ] 


By W. WATSON, D.Sc. 

Little attention has been paid for many years to the fungi occurring as para- 
sites on lichens in the British Isles, either by lichenologists or mycologists. 
During the last century a few fungal parasites on lichens were described 
by Mudd ( 1 86 1 ) , Lindsay (mostly in 1 869) and others, but it was not till this 
century that any serious attention was paid to them, A.L.Smithin igiogave 
an account of some in these Transactions {54a) ; in her first edition of British 
Lichens^ voL ii ( 1 9 1 1 ) about twenty parasitic fungi were mentioned, and in the 
second edition (19^26) the list was extended to about sixty, but many others 
were included as lichens in the general text. In the lists of Pyrenomycetes and 
Hyphomycetes published in these Transactions for 1940 and 1941 the names 
of some of our fungal parasites were given, some have been incidentally 
mentioned in my Lichenological Notes published in the Journal of Botany 
and elsewhere, but many British species have been entirely ignored. 

Amongst foreign authors during the latter part of the last century 
valuable contributions were made to the study of these lichen parasites by 
Rehm, Saccardo and Zopf, but the most thorough work must be attributed 
to the later studies of Olivier (1905-7), Vouaux (1912-14) and Keissler 
(1930), Die Flechtenparasiten of the , last-named refers to many British 
records and is of such an excellent standard that its arrangement and, 
nomenclature are usually followed even when they differ from those 
adopted by British mycologists. Other names in use or which have been 
used by British botanists are given so that any reference to a British plant 
can be traced. Prominence is given to British references and foreign ones 
have been largely ignored, often even when the nomenclature used depends 
on one, though this reference can be obtained through the references given. 
Some account of their distribution in Great Britain and Ireland has also 
been included. In most instances when no good description is readily 
available one has been added. 

Systematic arrangement (after Keissler) 


Pezizinae: Celidaceae, Helotiaceae, Mollisiaceae, Patellariaceae. 

Phacidiineae : Stictidaceae. 

Pyrenomycetes , ^ , ^ 

Perisporiales: Eurotiaceae, Perisporiaceae, Microthyriaceae. 


Dothideales: Phyllachoraceae. 




3o6 Transactions British Mycological Society 

Pyrenomycetes [continued) 


Sphaeriales : Sphaerellaceae. 

Pleosporae: Pleosporaceae. 

Fungi imperfecti 

Phycomycetes are not included, but occasionally some species may be 
found on lichens though they are not really parasitic on them, 

Keissler’s Flechtenparasiten includes some Myxobacteriales and Myxo- 
mycetes as lichen parasites and gives some British references for them. 

Myxobacteriales: Chondromyces aurantiacus Thaxt. in 2% 33; Stigmatella 
Berk. & Cke. in Introd. crypt Bot (1857), 313 ; Polycephalum Kalchb. & Cke. 
in 14, XIV (1884), 23; ? Stilbum rfytidiospora Berk. & Br. in J , Lim, Soc. 
(Bot.), XIV (1873), 96. 

Myxococcus fulvusjahn, in 23, 34 ; M. pyriformis A. L. Sm. in 20 ( 1 90 1 ) , 7 1 . 
Miss Smith’s plant was found at Llanwymawddwy, Merionethshire, in 
1899, on rabbit pellets and not on a lichen. 

Myxomycetes: Hymenobolina parasiticus Zuk. in Lister, Mon, Myc. 3rd ed. 
(1925), 1 81; Hymenobolus parasiticus Zuk. in Lister, Mon, Myc, 2nd ed. 
(1911), 262. This is given as parasitic on lichens, but no British locality is 
given for this habitat. 

Lister ella par adoxa‘}^n, in Lister, Mon, Myc, 2nd ed. (191 1), 262 ; 3rd ed. 
( 1 925) , 25 1 . This occurs on Cladonia podetia, but no British locality is given. 
Dicfymium melanospermum Macbride in Lister, Aloti, Myc, 2nd and 3rd ed. 

(1925)- . 

Craterium minutum Fr. in Lister, Mon. Myc. 3rd ed. (1925), 95. 

Badhamia capsulifera Berk, in 68, xxi (1852), 153; in Lister, Mon, Myc. 
3rd ed. (1925), 10; J?. hyalina in Berk. loc. cit.; in Lister loc. cit. 

B, utricularis Berk. loc. cit. ; Lister loc cit. 

B, versicolor Lister in 20, xxxix (1901), 81; Lister, Mon, Myc. 3rd ed. 
(1925). Occurs on bark and lichens in Aberdeenshii'e. 

In the following list fungi which have been included as lichen parasites 
but are not really so are usually enclosed in square brackets. A similar 
method has been employed for fungi which have been considered as lichens 
and for lichens which have been included in fungi. Lichens which appear 
to be parasitic occasionally occur but are rare. One of the most striking 
examples seen was noted near Llangollen where Biatora viridescens Fr. was 
growing over a Fomes. 




CoiMopliora laxa (Fr.) Quel.; Corticium Fr. 52, 626 (with description), is 
included in Keissler’s list. It does occur on lichens but is probably 
entirely saprophytic. 

British Fungi Parasitic on Lichens. W. Watson 307 

Corticium arachnoideum Berk. 52, 676; i, xni, 344; 41, i, 122 (with 
description); C. centrifugum (Lev.) Bres., Rhizoctonia Lev., Sporotrichum 
lichmicolum Berk._& Br. in 21, xrv (1872), 122, apud 23, 523. Occurs 
on corticicolous lichens, especially Xanthoria parietina, but also on many 
other lichens. 


[Agyrium rufum Fr. This fungus, which is given as a saprophyte on wood 
in 41, IV, 1 70, is included as a lichen, as it occasionally occurs asso- 
ciated with a lichen thallus, in 20 (1867), 257; 31, 392; 8, 10. The 
following localities are given for this supposed lichen: Knighton, v.c. 
43; Dolgelly, v.c. 48; Achrosagan, v.c. 98; Ballynahinch in I, v.c. 16 
and Killarney in I, v.c. 2.] 

Celidium lichenum (Somm.) Schroet., Dothidea Somm. in 23, 89; C. stic- 
iarum Tul. apud Linds. 68 (1866), 506; Sphaeria DN.; C. dubium Linds. 
69 (1866), 449; Homostegia Gke. 14 (1886), 66. Sticta pulmonacea var. 
pleurocarpa Ach. is a state of the lichen, Lobaria pulmonaria, in which the 
apothecia are abortive, tuberculose-diiform and brownish owing to 
the attacks of this fungus, 9, 272 ; 55, 1 16. This state has been noted on 
specimens from Bocconoc (2), Hafod (46), Gwm Bychan (48), Tros- 
sacks (87), Gawdor (96) and Appin (98). Lindsay also records the 
fungus as parasitic from v.c. 90. 

[C. squamaricolum Linds. 50 (1869), 142 is expunged by Keissler. 
According to Vouaux it is a sterile mycelium, formerly placed in 
Antennaria and now in Racodium^ 71.] 

C. varians (Nyl.) Arn. 41, iv, 10; 78, 62; Lichen^ Hall, Brit FL; Davies in 
68 (1794); Arthonia Nyl. in 57, 240; Arthonia NyL 1856 ex Leighton 
in I (1856)5 330; Lecidea epip^ta Stirt. 21 (1874), Arthonia 
galactinaria Leight. p.p. 31, 426; Conida Vouaux, 57, 390. Description 
in 56, 2 1 8” 1 9. Parasitic on the thallus oiLecanora rupicola [glaucoma) 
with records from the following vice-counties: G, i, 2, 4, 40, 42, 45, 
48, 49, 52, 62, 69, 71, 9I5 92, g8, 100; Ireland, 3, 6, 12, 16, 21, 27, 
35, 38. Form parasemoides (Nyl.) Arn. 23, 96; Arthonia parasemoides 
Nyl, in 42, 252. Differs from the type by the paraphyses being almost 
absent and the spores 2-septate, the middle cell somewhat greater. 
Usually on the apothecia of Lecidea parasema. From Gunwalloe, 
v.c. I, Longmynd, v.c. 40, Cleveland, v.c. 62, and Lerwick, v.c. 112. 

C. varimn (TuL) Massal. 23; 71; 57, 390; Phacopsis Tul. in 23; Lecidea 
glaucomaria Nyl. 1852 in Leighton 69 (1878), 238; 31, $8g; Leciographa 
A. L. Sm. 56, 186 and 57, 202 (which see for description) . Note that 
Lecidea glaucomaria Nyl. of 1852 corresponds with this fungus whilst the 
Arthonia glaucomaria Nyl. of 1856 corresponds with Celidium varians. 
Both occur on the same species of Lecanora^ but C, varium is found 


3 o 8 Transactions British Mycological Society 

on the thallus whilst C. varians occurs on the apothecia. As the 
spores are 3-septate and brown the species goes under the subgenus 
Celidiopsis Sacc. Recorded from the following v.c.: 5, 45, 48, 71, 90, 
96 and 1} 16, 21. 

[Conida abroAallus Linds, is given by Arnold in Flora (1878), 104, by the 
name alone and is discarded by Keissler in 23, 86.] 

Conida demens (Tul.) Massal., Phacopsis Tul. 23, 72 ; Arthonia punctella Nyl, 
in 5 (i 859), 553 ; 42, 252 ; 57, 241 ; A. subvarians Nyl. in 56, 2 19 ; 57, 241 ; 
A. galactinaria Leight. p.p. 31, 426; ? Microthelia subfuscicola Linds. 50 
( 1 87 1 ) , 39 ; Conida punctella Arn. in 56, 346 and 57, 390. Lindsay records 
it as occurring on the thallus of various Lecanoras, but it also occurs on 
man y other lichens. Both C. punctella Arn. (with somewhat broader 
spores and hyaline hypothecium) and C. galactinaria Arn. (with spores 
becoming brownish) are included by Keissler in this variable plant. 
Recorded from v.c. 5, 14, 33, 48, 52, 71; I, 5, 16, 21, 27. 

C. epiphorbia (Stirt.) Vouaux, 71; 57, 390; Leddea Stirt. 14 (1873), 108; 
31, 388; 56, 104; 57, iii; Karschia Zopf in 51, 378. On Solorina bispora 
Ben Lawers. Keissler considers that the species is insufficiently 

C. fuscopurpurea (Tul.) Vouaux in 57, 390; Celidium Tul. in Linds. 50 
(1869), 141; Arthonia peltigerea Th. Fr., Scutula Rehm in 57, 241 and 
389 ; Homostegia pelvetii (Hepp) Linds, in 4, 206 with further references ; 
Conida pelvetii Arn. is included in 23, 77 as a synonym. On thallus of 
Peltigera and Solorina, Corriemulzie, v.c. 92. Form stereocaulina (Ohl.) 
Keissl. Arthonia nephromaria var. Ohl. differs from the type by the 
hymenium becoming red straightway with iodine and in its habitat 
on Stereocaulon or occasionally on Physda. 


Pezizella epithallina (Phil. & Plow.) Sacc., Peziza Ph. & PI. 14 (1887), 
24; Mollisia Phil. 47, 173; Mollisiella Bond, in 51, 377; Peziza miliaris 
Wallr. in 23, 108; 47, 458. Collected by Plowright at King’s Lynn. 


Orbilia coccinella (Somm.) Karst. 51, 372; 41, rv, 144; Peziza Somm., 
Calloria Phil. 47, 328-9. Usually on dead wood but sec. Rehm also on 
crustaceous lichens where it is probably saprophytic rather than 


Nesolecbia associata (Th. Fr.) Sacc. & D. Sacc., Th. Fr. in 23, 

igSj i. leptostigma Nyl. in 57, 52; Nesolechia leptostigma Sacc. as syno- 
nym in 71 ( 1913), 415 and 23. A. L. Smith puts this in the Biatora 
section, gives references to Crombie and Leighton, states that it occurs 
on Ochrolechia tartarea on a mica-schistose boulder on Ben Lawers and 
suggests, as Nylander thought the thickish thallus was not proper, 
that the apothecia belong to a fungus parasitic on a sterile thallus. 

British Fungi Parasitic on Lichens. W. Watson 309 

N. cetraricola (Linds.) Am. in 57, 389; Lecidea Linds. 50 (1869), 57 and 
(1871)5 35; 68 (1871)5 364 and PL 48, On Cetmria islaniica zxA C. jmi- 
perina at Braemarj v.c. 92. The description is insufficient for the species 
to be included in Keissler’s key. 

N« cladoniaria (Nyl.) Arn. in 57, iii and 389; Lecidea Nyl. in 8, 94; 30, 
358; 5^5 ^^41 32^5 388; Abrothallus cladoniamm Linds. 39, xxv, 546; 
? Ab. moorei Linds. 39 (1869)5 554. Parasitic on Ck£/c??2k5 Kelly’s Glen 
near Dublin. 

?N. insita Vouaux, 71; 575 389; Lecidea Stirt. in 63 (1879)5 17; 31, 545 
and 57, 1 12. Parasitic on Peltigera aphthosa and collected by Stirton at 
Greag-na-Lochan, v.c. 88. It is listed as a lichen in British licheno- 
logical works and may be a form of Biatora geopham (Nyl.) Fr. The 
thallus of the latter is often so indistinct that the position of the plant 
is rather doubtful. A plant collected in 1928 on marly soil at Orchard 
Portman, v.c. 5, was considered at that time to be a fungus, but 
recently a re-examination showed that it was identical with a specimen 
of the Little Bowden plant described as Lecidea pleiospora A. L. Sm. in 
2O5 XLIX5 41 and 563 3525 and later placed under L. geophana Nyl. in 
57 . 5 ^- 

[?N. inttunescens Magn.5 Lecidea intumescens (Flot.) Nyl.j L. badia var. 
intumescens Flot.j L. insularis Nyl. in 565 94 and 57, 104. Magnusson 
suggested that this was a JVesolechia^ but3 though it is apparently 
parasitic on Lecanoras and other lichens and partially destroys the 
thallus over which it grows, it is considered to be a true lichen. The 
designation Lecidea insularis is adopted by A. L. Smith, as Nylander’s 
insularis dates from 1852 whilst intumescens was not used as a specific 
name till 1876.] 

N* lichenacola (A. L. Sm. & Ramsb.) Keissl. 23, 140; Discocera A. L. Sm. 
& Ramsb. in 59 (1917), 48; 77, 93. Found in 1915 and 1916 on 
liclien-thailus on stone at three different places near Taunton, Britty 
Common, Feltham and Treborough. The lichen was sterile in every 
locality and was not determinable. Keissler states that it is ^ eine sehr 
merkwurdige Art’. 

N. neglecta Vain. 70, iv, 418-19; 74, vm, 152. ^Lecidea neglecta Nyl.’ in 
Crombie, lo, xiii, 14 1; Leight. 31, 276; A. L. Sm. ‘56, 96 and 57, 
1065 refers to the thallus on which the apothecia of the fungus are 
situated. This is given by Lynge in 70, iv, 419 as Crocynia neglecta 
Hue; but also see 74, tom. cit. 

N. ©xyspora (TuL) MassaL 71; 23; 57, iii and 389; 78, 62; Abrothallus 
TuL in 38 (1857), 37; 42, 225; Lecidea Nyl. in 56, 103; 57, in; 8, 92; 
3 ®. 3575 3^? 384* Lecidea obscuroides Linds, in 69, xxn (1859), 247 and 
PL XIII, figs. 36-8. This was described from a specimen collected 
by Carroll at Dunscombe Wood, Cork. Lindsay’s description agrees 
quite well for jV*. oxyspora^ though the spore measurements are rather 
smaller. Parasitic on various Parmelias and recorded from v.c. 1-3, 
35. 37. 48, 49. 64, 72, 81, 87-92, 97, 98, lOi, 104, no; Ireland I, 
3» 16. 

3 ^^ Transactions British Mycological Society 

N. piincta Massal. in 23, 126. A specimen from Redhead, Forfarshire, 

: was considered by Lamb to belong here probably, but the hymenium 

is not coloured by iodine. N. inquinans MassaL, if synonymous, appears 
to have priority. 

N. viteliinaria (Nyl.) Rehm, 71; 23; 57, 389; 78, 62; Lecidea Nyl. 42, 212; 

' % 78; 30, 355; 31, 384; 56, 60; 57, no. Parasitic on the thallus of 

' Candelariella mtellina and also on other crustaceous lichens. Recorded 

from v.c. I, 35, 37, 40, 62, 88-90. 

Var. supersparsa (Nyl.) Keissl. Lecidea Nyl. in 23, 133, has spores which 
are more or less rhomboid and acute at their apices. A specimen 
collected by Miss Duncan from Lundie Crags has been seen by me. 
Scutula cristata (height.) Sacc. & D. Sacc. 51, 379; 74, vi, 71; 77, 93; 
Lecidea height, 31, 385; Biatorina A. L. Sm. 57, 143; Catillaria Arn. 
(see 74, VI, 71). On the thallus of Lecanora subcarnea at Barmouth, but 
on other species at the Lizard and Arbroath. 

S. epiblastematica (Wallr.) Rehm, Peziza Wallr. 23, 149; Biatorina 
A. L. Sm. 57, 144; 77, 38; ? Lichen glebulosus Smith, Eng, Bot. 1809; 
Lecidea wallrothii Nyl. 20 (1874)^ 148; 31, 388; non 56, 29 and 57, 19; 
L, heerii Hepp. in 20 (1882), 274. On the thallus of Peltigera and 
Solorina on Craig Calliach, Glen Lyon and Ben Lawers. 

S. epicladoiiia (Nyl.) Sacc., Lecidea NyL, Conida Vouaux in 23, 154; 
Biatorina Arn. in 74, vi, 72, which see for description. On the 
squamules of Cladonia pyxidata var. pocilla at Portland, v.c. 9. 

S* episema (Nyl.) Zopf in 23, 160; Lecidea Nyl. in 20 (1867), 257; 8, 78; 
31, 385; Biatorina A. L. Sm. 57, 143; 77, 38; Catillaria Oliv. in 74, ix, 
36; Biatorina supernula A. L. Sm. 57, 143; Lecidea Nyl. in 31, 389. On 
the thallus of Aspicilia calcarea and fairly common. Recorded from 
v.c. 5, 6, 33, 37, 41, 48-50, 88, 103; Ireland i, 5, 7, 8, 10, 16, 17, 29. 
[S. peltigerea Rehm, Arthonia Th. Fr. in A. L. Sm. 57, 241 is included 
by Keissler under Conida fus€Opurpuread\ 

S, stereocaulomm Krb. in Vouaux, 71 (1913); Biatorina Th, Fr. in 57, 
143, with description; Lecidea Nyl. in Crombie, 14, xxn, ii. On the 
phyllocladia of Stereocaulon. No locality is given by A. L. Smith, but I 
have recently seen this parasite on a specimen of S, denudatum from 
Achnacashen, v.c. 106. 

Mycobilimbia endocarpicola (Linds.) Vouaux in 57, 389; Lecidea Linds. 
39 (1869), 547 and 50 (1869), 136. This is incompletely described sec. 
Keissler. ^ Found by Carroll in England on Dermatocarpon liepaticum 
and may have to go in the genus Celidium\ 23, 171. 

M* tdUiasii (Hepp) Rehm, 23. This has recently been noticed on a 
specimen of Peltigera aphthosa collected in 1929 from near the summit 
of Ben Lawers. 

[M. obscurata (Somm.) Lecidea sphaeroides yzx, obscurata Somm. 

23, 1 71. This is included by Vouaux as a lichen parasite. Though it 
may grow on the thallus of Peltigera^ Keissler considers it to be a true 
lichen. It is given in 83 by Zahl. as Bacidia obscurata and by A, L. Sm. 

British Fungi Parasitic on Lichens. W. Watson 31 1 

in 56, 143 and 57, 156 as Bilimbia sabuletorum var. obscurata. Another 
species, Bilimbia effusa Auersw. (in 77, 59), is not a lichen parasite as 
some authors state but a true lichen, 23, 172,] 

[Mycobaciiiia arenicola (NyL) Sacc. in 51, 378; Lecidea Nyl. in 31, 386; 
Raphiospora Mudd, 42, 186. This is given by Vouaux as a lichen 
parasite, 71 (1914), 140, but is usually considered to be a lichen in 
which the thallus is obsolete, the apothecia often growing over the 
thallus of Baeomyces. It is given in 56, 165 and 57, 180 as Bacidia flavo- 
virescens var. arenicola A. L. Sm.] 

[M. flavovirescens (Dicks.) Rehm; Lichen Dicks, in 82; Lecidea Borr. in 
61; Raphiospora Krb. in 42, 186; Lecidea citrinella Ach. in 8, 94 and 31, 
366 is given by Vouaux and some other authors as a lichen parasite. 
It is a true lichen sec, Tobler and many other lichenologists. Though 
it may grow over the thallus of Baeomyces it often occurs where no 
Baeomyces is present. It is listed by A. L. Sm. (57, 179) as Bacidia 
flavovirescens Anzi, and is not uncommon in subalpine regions.] 

M. vermifera Vouaux is given by Keissler under Spilomela. 

M. plumbina Vouaux is given by Keissler under Lahmia, 

Lahmia fuistingii Krb. in 23, 175. Spores at least 8-celled, 20-40 (-70) x 
2-2 *5^6, to 8 -nae, more or less parallel above, hyaline, acicular. Ascus 
clavate, 90-1 00 x 1 5-1 7 /i,, above somewhat acute. Paraphyses septate. 
Hypothecium almost colourless. On Baeomyces rufus, Britty Common^ 
near Taunton, Somerset. 

L. plumbina Keissl. 23, 176; Leciographa Anzi in 56, 186; 57, 202; 51, 378; 
Mycobacidia Vouaux (1914) in 57, 390. On Parmeliella plumbea in 
Borrowdale, v.c. 70 and Lowther Park, v.c. 69. Description in 56, 186. 

Mycomelaspilea leciographoides Keissl. 23, 218; Melaspilea Vouaux 
71 (1913)5 472; 83, n, 248. Apothecia mostly 2-3 or even to 6 aggre- 
gating or nearly so, sessile, at fost punctiform but soon lengthened and 
lirelliform, with thick margin. Ascus 75-80 x 23-28 ju. Spores 8-nae, 
biseriate, strongly constricted between the longer upper hyaline to 
brownish cell and the lower smaller cell, 19-22 x io-12/x. Epithecium 
and hypothecium brownish. Iodine colours the contents of ascus and 
paraphyses blue then reddish brown, the ascus wall becoming blue 
and then yellow. On Verrucaria, Beetham, v.c. 69. 

KarscMa adjuucta (Th. Fr.) Arn.; Buellia Th. Fr. 23; 71; 74, vii, 332, 
which see for description. It is incompletely described by Th. Fr. sec, 
Keissler. The coloration of the hymenium with iodine is blue, whereas 
Vouaux gives it as negative. Known from v.c. 41 on Lecanora expallens 
and L, conizaeoides^ and from v.c, 58 on the latter host. 

K« advenula (Leight.) Zopf, 51, 378; Lecidea height. 68 (1876), 146; 31, 
338; Buellia A. L. Sm, 57, 200; 74, vin, 155. Keissler apparently 
followed A. L. Smith in giving the spores as 4-nae. They are given as 
8-nae by Leight. 31, 388, and the type specimen at Kew seldom had 
less than 8 spores in the ascus when it was examined by me. On the 
thBllm of Pertusaria, Llanbedrog, v.c. 49, Appin, v.c. 98 and Garryna- 
hine, v.c. no. In Ireland it was collected by Larbalestier in Conne- 
mara (I. 16) and in Mayo (I. 27) ; 25, 371. As an instance of confusion 

312 Transactions British Mycological Society 

between lichens and lichen parasites Lecidea epipsila NyL {Buellia Oliv.) 
is mentioned by A. L. Smith in 56, 185 as being allied to Buellia 
[Karschia) advenula^ both occurring on the same species of Pertusaria, 
Keissler, 23, 201, states that it is a lichen with a thallus of its own, even 
though this grows over the thalli of various crustaceous lichens, 
especially Pertusaria wulfeni form rupicola ( = P. sulphurea Schaer.). 

K« athallinia (Mull.) Vouaux; Buellia Mull. 23, 183; Lecidea particularis 
NyL in 31, 386; Buellia A. L. Sm. 56, 184; 57, 200; 25, 370. On the 
thallus of Baeomyces at Kylemore and Dungannon in Ireland and 
recently collected by Sowter at Sandsend in Yorkshire, v.c. 62. 

K. destructans TobL, which occurs on Chaenotheca^ is mentioned in 58, 
265, but no British records are known to me. 

[K. lignyota (Fr.) Sacc. in 51, 378; 41, iv, 99; 75, iv, 208; Patellaria Fr. in 
47? 360 was included amongst the lichens by Leighton as Artkonia 
melaspermella NyL in 31, 416.] 

K. saxatilis (Rabh.) Rehm, Trachylia Rabh., Calicium Schaer. in 23, 
191. Calicium saxatile Schaer. is given by A. L. Smith (57, 196) as a 
synonym of Buellia saxatilis Krb., and this is retained as a lichen by all 
British authors. The lichen parasite has not yet been recorded for the 
British Isles, though it probably occurs. 

[K* scabrosa Rehm in 23 and 71; Buellia scabrosa form athallina A. L. Sm. 
57, 196. This is not a true parasite, but the lichen Buellia scabrosa Krb., 
growing over Baeomyces rufusi\ 

K. talcophila Krb. in 23, 189; Lecidea ferdinandezii Gromb. 21 (1877), 223. 
Crombie’s reference is to a foreign specimen. 

K. thalloides (Hepp) Rehm, 23, 199. A parasite on Pilophorus from 
Killarney agrees with this species except for the^ host which usually 
is Physcia, 

Abrothallas microspermas TuL 23, 213; 71 (1913), 468; A, smithiivdx. 
microspermus Linds. 50 (1857), 34 68 (1866), 507. On Stictina 

fuliginosa at Tavistock (specimen in herb. Kew.). 

A. moorei Linds. 39 (1869), 554. On Cladonia uncialis and G. bellidiflora. 
Vouaux considers this as a synonym of Phoma uncialicolay but Keissler 
does not think Vouaux’s theory possible as Lindsay gives to his species 
an ascus with spores. He considers that the species is better eliminated, 
though it may refer to JSfesoleckia cladoniaria. 

A. parmeHarum (Somm.) NyL in 23; 71; 78, 62; Lecidea Somm. in 8, 92-; 
30, 357; 3I5 386; Buellia Oliv. in 56, 183 and 57, 199; Lichen parasiticus 
Smith, Eng, Bot, (1808, and so prior to starting-point of fungal nomen- 
clature) ; A, smithii TuL in i (1852), 1 13 and 50 (1857), 34; Mudd, 42, 
224; Lecidea apud Linds, in 69 (1859), 292 ; Phymatopsis dubia Linds. 69 
(1866), 442; Abrothallus parmeliarum var. ater Linds. 38 (1857), 34; 
A^^^ is a nomen nudum sec, 23, 207, but was described 

^ zs A, welwitschii TuL Occurs on foUaceous and fruti- 

culose lichens and is common. It has been recorded from 27 English, 
13 Scottish and 8 Irish botanical counties. When it occurs on 'Usnea 
as it does' at Llanbedr, v.c. 48, it has been styled Abrothallus usneae 
Rabh., A, form usneae by Rehm and Vouaux and Buellia:,. 

British Fungi Parasitic on Lichens. W. Watson 313 

usneal by Jatta (74, vii, 332). Keissler includes this and some other 
creations with the type. Form chrysantheus (Stein.) Arn. has the 
apothecial disk remaining greenish golden pruinose right up to an 
old stagCj whereas in the type the pruina is some other colour and 
is lost much earlier. Leighton’s no. igi, 2% is probably this form 
sec. Keissler^ 235 212. 

Leciographa centrifnga (Mass.) Rehm, in 23, 240, probably occurs in 
our islands, but the only record is a doubtful one. 

L. inspersa (Tul.) Rehm, in 23, 226; 71 (1913), 476; 26, ii, 165; Lecidea 
Flk. 1819, so before starting-point for fungal nomenclature; Dactylo- 
spora Mudd, 42, 224; ? Calicium stigonellum Salwey in 66 (1846), 204, 
non Schaer.; 2% no. 183; Lecidea zwackhii Cromb. i© (1876), 362 non 
MassaL; Leciographa parasitica Massal. in 56, 185; 57, 201; 77, 61 sed 
non Massal. (for description of L. inspersa see this). On the thallus of 
Ochrolechia parella^ Pertusaria species, etc. Not uncommon and recorded 
from the following botanical v.c. : C, 1-3, 5, ii, 13, 14, 19, 23, 26, 33, 
35*"8> 44 ^ 47 ~ 5 ^? 55^ 60, 62, 90, 92; Ireland, 2, 4-6, 15, 16, 20, 21, 35, 
38. 39- 

L. parasitica Massal. in 56 and 57 does not refer to Massalongo’s plant 
but to L. inspersa. It occurs in our islands and the paucity of 
records is probably due to the confusion of the two species. It differs 
from L. inspersa as follows: epithecium and hypothecium brown, 
hymenial gelatine blue then wine-red with iodine, spores larger, 

L. parellaria (Nyl.) Sacc. & D. Sacc. in 71 (1913)^ 486 and 23, 236; 
Lecidea NyL in 10 (1876), 362; 31, 387; Leciographa parasitica var. 
parellaria A. L. Sm. 56, 186; 57, 201.' On Ochrolechia parella^ Fishguard, 
v.c. 45 leg. Leighton, near Conway, v.c. 49 in Herb. Brit. Mus., 
Connemara, I. 16 leg. Larbalestier. 

L. lamyi (Rich.) Sacc. & D. Sacc., Lecidea Rich. 71 (1913)5 483 and 23, 
230. / . specimen from Arbroath on Lecanora rupicola seems referable to 
this species. Apothecia in groups of 3-10 with dark discs, o- 1 5-0*4 mm. 
Ascus may have 8 spores but often only 6 or 4. Spores 3-septate, 
without oil-drops, smooth-walled, elliptical, obtuse at both ends, 
hyaline at first but finally brown, 16-22x6-9^6. Hymenium with 
iodine blue then wine-red. Hypothecium dark brown. On Lecanoras. 

[L. scapanaria (Carrington) A. L. Sm. 57, 202; Lecidea Carr, in 66 
(1863), 382 and 41 1 ; 8, 87; 31, 387, is not a lichen parasite but is 
a fungus on a liverwort. L. persimilis Nyl. in 14 (1873), 71; 3I5 389 
is included by A. L. Smith. Recorded from vice-counties 88, 89; 
Ireland 2, 16.] 

[L. zwacMm Vouaux, Massal. 71 (1913)5 480; 23, 229. Lecidea 

zwackhii is by Crombie in 20 (1876), 362 as collected by 

Joshua and parasitic on the thallus of Pertusaria aX Chedworth in 
v.c. 33, but it is not Massalongo’s plant. Sec. A. L. Smith, Crombie’s 
record probably refers to her Leciographa parasitica {^L. inspersa 

314 Transactions British Mycological Society 

Spilomela vermifera (Leight.) Keissl. 23, 221 ; _ Melaspilea Leight. 68 
(1876), 146; 31, 437; 57, 251; Scutularia Sacc. in 51, 378-9; M. vermi- 
Jormis Leight. in 14 (1875), sec. 83 (1923), 284; Mycobacidia vermifera 
Vouaux 71 (1914), 141. On thallus and apothecia of Pertusaria 
globulifera on oak, Trefriw, v.c. 49. 

[Biatorella campestris Th. Fr. in 56, 353; 57, 115. This is considered 
by lichenologists to be a lichen, but the manner in which the apothecia 
develop (see 74, vii, 328) is very similar to that of a parasitic fungus.] 
[B. difformis (Fr.) Wain., B. fossarum (Duf.) Rehm, B. moriformis (Ach.) 
Th. Fr., B. pinicola (Massal.) Th. Fr. and B. resinae (Fr.) Mudd 
are given by Ramsbottom in 51, 379, amongst the fungi, but most 
■ lichenologists place them amongst the lichens; 8, 75-6; 31, 382-3; 
42, 191 ; 57, 1 15-17, etc- Massee in 41, iv, 95-6 includes B. resinae and 
B. pinicola amongst the fungi. Sec. A. L. Smith the terminology 
B. fossarum (Duf.) Th. Fr.; B.' resinae (Fr.) Th. Fr. is correct.] 
[Sphinctrina gelasinata (Wahl.) Zahl., Lichen Wahl, in 83, i, 581; 
Sphinctrina turbinata Fr. in 55, 3 and S', microcephala Nyl. in 9, 84; 
S. tubaeformis Massal. in 55, 3 are lichens parasitic on Pertusarias and 
have been given by Rehm and other authors as fungal parasites; in 
fact, Rehm considers the genus as a fungal one.] 

[Stenocybe major Nyl., S. euspora Anzi in 23, 643 and other species of the 
genus are given as fungal parasites on Tkelotrema, etc., but they are 
generally considered as lichens incidentally growing over other 
lichens, 9, 97 - 8 ; 55, 19-20; 31, 45; 42, 256.] 

[S. bryopMa Wats. 74, i, 130; n, 109; iv, 74; v, 265. Collected near 
Llanberis on Sphaerophorus melanocarpus [compressus) and neighbouring 
liverworts. Entirely independent research by Abbe Grelet on a 
specimen, found on a liverwort collected by S. M. Macvicar in west 
Inverness during 1903, resulted in the description of Stenocybe major 
var. macvicari (12) which was included amongst the fungi. The author 
later acknowledged the priority of the name bryopkUa.} 



Stegia^ vermicularis (Arn.) KeissL, Mesolechia Arn. z% 255. This and 
other parasites on Cerania vermkularis have been described as 'fruits? of 
the host. The apothecia form galls on the thallus, are pale, flat or 
convex, immarginate, loo/z- diam. The asci are cylindric or cylindrico- 
clavate, about 70^6 long and about broad in the broadest part, 
with 8 simple, hyaline, ± uniseriate, elliptical spores about 5-1 1 x 3-5 /r. 
The paraphyses are thread-like, discrete, simple, rounded at apices; 

absent, thecium hyaline and hypothecium yellowish or 
pale brownish, I — . On Cader Idris leg. Watson apud Keissler in Ann. 
Jsfaturk Mus. Wie% xxxvin (1925), 163. 

[Xylographa* The following species are included as fungi by Rams- 
bottom in 51, 380. They are included as lichens by A. L. Smith (57, 
245-7) and other British lichenologists.] 

British Fungi Parasitic on Lichens. W. Watson 315 

[X. parallela (Ach.) Fr. in 41, iv, 64; 8, 95; 31, 391; Stictis Ft, in. 61, 
213; X. abietina (Pers.) Zahl. 83 (1923).] 

[X* laricicola NyL in so, in, 128; 31, 391.] 

[X. scaphoidea Stirt. in 14, iii, 36; 31, 391; X parallela var. pallens ^Hil, 
in 57, 246.] 

[X. spilomatica (Anzi) Th. Fr. Ag^rium Anzi in Comm. Soc. Critt. ItaL 11, 
20; ? X. minutula Krb. Par. 276.] 

[Odontotrenaa. This genus was included in the lichens by Crombie % 10. 
The species put under this genus in Leighton's Lichen Flora occur on 
wood, have very indistinct thalli and are better included in fungi. The 
following have been given as British plants.] 

[O. firmatum NyL in 20 (1882), 276.] 

[O. longiiis NyL in 31, 389 and 51, 380.] 

[O. majas height. 31, 390. Spores 19-21 x 7'5~8-5/x.] 

[O* miniis NyL in 31, 390; 20 (1882), 276. Spores 12-14x5/^1.] 
[Melittosporiani lichenicolum (Mont.) Mass. 41, iv, 88; 51, 380. This 
name has been given to the form of Diploschistes scruposus which occurs 
on the squamules of Cladonia and therefore considered by Massee and 
others to be a lichen parasite. It is, however, a true lichen and many 
intermediate states between the typical lichen with a proper thallus 
and the athalline form on the Cladonia squamules may be traced. It 
is given as Urceolaria scruposa var. bryophila form lichenicola (Rich.) 
Harm, in Harmand, Lich. France, 1150; by Crombie in 14 (1891), 60 
as Lecanora scruposa subsp. bryophila form ecrustacea; by A. L. Smith, 55, 
383, under Diploschistea bryophilus Zahl. and in Zahl. Cat. Univ. as 
D. scruposus var. parasiticus (Somm.) Zahl. 73, 298; 74, x, 147; 77, 66. 
Sometimes the spores in the athalline form are less developed than 
they are in neighbouring typical plants.] 

[ScMzoxyloJi corticola (Fr.) NyL, Coniangium Fr. is recorded as a 
lichen on old oaks near Lyndhurst by Leighton in 31, 350. This 
species is not given by Massee, who states in 41, iv, 69, that Schizoxylon 
Pers. is not that of Leighton.] 

[MelaspUea patersom Stirt. in 54/ iv (1887), 29; 31, 437; 56, 225. It 
was collected from dead bark at Ben Brecht, v.c. 98, by Dr Paterson 
and is stated to be a discomycetous fungus, Schizoxylon species, in 
57, 248.] 




Eiirotiiiiii herbariomin Link, ex Fr. For: references see 4, i34* This is 
given in Keissler's list. It is not uncommon on imperfectly dried 
lichens, but rarely occurs on living specimens and is probably better 
considered as a saprophyte. 

3 i 6 Transactions British Mycological Society 


Orbicula tartaricola (Nyl.) Cke. in 6, 926; 4, 137; 23, 267; Sphaeria 
Nyl. apud Leighton in i (1876), 408. On thallus of Ochrolechia 
tartarea, Cader Idris, Hengwrt, near Dolgelly. 

[Strigula babingtonii Berk., Eng. Bot. Suppl. t. 2957 (1849),- 28, 70; 
78, 62; is included by Leighton, 31, 498 and Mudd, 42, 309, as a 
lichen. Zahlbruckner in 83 states that it is a fungus. Both Hariot 
(J. Bot. Paris, HI, 285) and MuU.-Arg. {Flora, Lxxm, 200-1) had 
previously excluded it from the lichens as it had no algal cells. It is 
probably a perisporiale, but there is no reference to it in 4. Records 
from Sussex, Gloucestershire, Cambridge, Flint, Leicestershire, York- 
shire and Durham have been given. Description in 31 and 42.] 


Phragmothyrium cetrariicolum (Nyl.) Keissl. 23, 273; Sphaeria Nyl. 
in 14 (1874), 68; Metasphaeria Sacc. in 4, 178; Psilosphaeria Cke. in 14 
and 62. On Cetraria, Braemar, in Scotland. 


[Broomella leptogicola (Cke. & Massee) Sacc. 23, 286; in 4, 176; 
Hypocrea Cke. & Mass, in 14, xix, 86 and described from a specimen 
found on the thallus of a Leptogium on Robinia at Kew. Petch (45) 
gave a description of it, regarded it as the fructification of a lichen 
and discarded the name. Von Hohnel {Ann. Myc. xv (1917), 237) 
regarded it as insufficiently described and suggested that it might be 
a Tatesula. In the same journal (1918), 93, he gave the generic name 
of Keissleria to it.] 

[Calonectria leightonii (Berk. & Br.) Sacc. ; Nectria Berk. & Br. This 
name was given to a plant collected from Yorkshire by Leighton on 
a larch. Petch (45) states that the British specimen is a lichen and 
should certainly not be included with JV. leightonii if the .Cuban 
specimen is taken as type. He also rejects the plant in 46, 300; 4, 195.] 

N. epicallopisma (Arn.) Sacc., Cercidiospora Arn. 23, 279, occurs on 
Placodium and has been doubtfully reported from Britain. 

Nectria indigens Rehm, Secoliga Arn. 23, 278. A specimen with a 
pale red apothecium which was collected at Dartington, Devon, was 
placed here by Keissler. 

N. lecanodes Ces. 23, 276; Dialoneciria Berk, in 14, xii, no; Lasionectria 
(Sacc.) Cke., Petch in 67 (xxi, 253) and 4, 198, is a semi-saprophyte 
on Peltigera. Recorded from King’s Lynn and North Wootton, Norfolk, 
by Plowright and from Inverary by Boyd. 

N. insidiosa Sacc.; Hymenobia NyL; Psora aporea Nyl., is given in 23, 279, 
as occurring on Lecideas in England. 

Nectriella robergei (Mont. & Desm.) Weese. Nectria & Desm.; 

N. lichenicola 'Kchm', N. peltigerae Phil. & Plowr, 4, 200; 23, 281; 46, 
270. Description in 46, 270. Occurs chiefly on Peltigera thalli. 
Recorded by Plowright from the King’s Lynn district and from 
Forden by Vize. 

British Fungi Parasitic on Lichens. W. Watson 317 

Ciliomyces oropensis Hohn. Sphaeria qffinis Grev. in 13; Mectria affinis 
Cke. in 62, sec. Fetch (46), who puts the plant as Parametria affinis 
(Grev.) Sacc., and states that it was found on Ephebe lanata at Appin 
by Carmichael. Description in 45, 201 and 46, 282. Other references 
given in 4, 201. 

[Tlielocarpon epithallmum Leight. 31, 439; 56, 345, is parasitic on the 
thallus of Baeomyces rufus at Stiperstones, Shropshire. It was included 
amongst the fungi by Rehm and Vouaux, the former regarding it as 
one of the Hypocreales, but Fetch in 46 does not mention it. Zahl- 
bruckner and other lichenolo^sts, on account of the well-defined 
gonidial sheath of the perithecia, place it with the lichens, 23, 295.] 



Homostegia pelvetii (Hepp) Linds, in 39 (1867), 450, is referred by 
Keissler to Conida fuscopurpurea, 23, 303 and 4, 206. 

H. piggotii (Berk. & Br.) Karst. 4, 206 and 23, 300; Dothidea Berk. & 
Br. in i (1852), 385; ? D. homostegia Linds, in (q, xxvn, 365 sec. 23, 
301 . On the thallus of Parmelia or other foliaceous lichen, Beddgelert, 
Capil Gurig and Barmouth. 


Adelococcus alpestris Theiss. & Syd. in 23, 309. The Ingleborough 
parasite mentioned in 75, m, 206, may be this, but description too 
incomplete and the host was not Acarospora. 

MuUerella atricola (Linds.) Sacc. in 23, 319; Microtkelia Linds. 39 
(1869), 554. Apothecia scattered, superficial, globose with somewhat 
flattened apices, small and dark. Ascus 50X with many spores 
which are globose, dark brown and about 2 [x diam. Keissler does not 
consider that the position of this plant is fully established. It was 
found on the thallus of Lecanora atra at Derryquin, Kerry. 

M. haplotella (Nyl.) Arn. in 56, 345; Endococcus Nyl. in 5 (1868), 10 1; 
Verrucaria Leight. in 31, 495, is placed by Keissler under Mullerella- 
polyspora, but has been considered distinct by some authors because 
the apothecia are darker coloured at maturity, 57, 386. 

M. opegrapMcola (Linds.) Zopf. Apparently given as a Microihdia 
collected by Carroll on Opegrapha atra at Aghada, Cork, by Linds, in 
39 (1869), 543 and 549. Ferithecia black, scattered, almost superficial, 
globose and flattened above, small. Ascus many-spored. Spores 
simple, round and brown sec. Keissler, 17 x B-^fx (23, 319). Decidedly 

requires further examination before it can be accepted. 

M. polyspora Hepp (1862) in 23, 315 (including M. haplotella) ; A. L.Srmth 
in 67 (1908), 1 16; 4, 153. It occvacs on Arthonia, Bacidia, Lecidea, 
Opegrapha and Verrucaria, and has been recorded from Jersey, Ben 
Lawers and Killarney. Ferithecia scattered or + grouped, at first 

3 i 8 Transactions British Mycological Society 

immersed but later emergent, about 1 20-1 70 /x diam. Asci ± cylin- 
drical-obovate, 150 x 50-60 /x many-spored. Spore simple, about 
X 2-3*5 brown, and sometimes with two guttulae. On 

various crustaceous lichens. 

Rhagadostoma licliemcolum (DN.) KeissL 23, 320; Bertia DN. 4, 157; 
Sphaeria Karst, in 14 (1873), 156; Psilosphaeria Gke. & Plow. 14 
(1879), 85; 62, 389; ? Microthelia solorinaria Linds, in 50 (1869), 349; 
Diplodina solorinaria Vouaux, 71. Grombie, in 8, 280, states that the 
parasitic fungus, Sphaeria lichenicola DN., is occasionally seen on the 
thallus of Solorina crocea^ a rare lichen, though locally abundant on 
the summits of some of the Grampians. 


[Myriangium duriaei Mont. & Berk. 4, 207; 75, i ; 77, 93. This has been 
included in lichens by Leighton (31, 37), Grombie (9, 15) and Taylor 
(as Collema glomerulosum Tayl.) in ii, 2, 108.] 



Guignardia corniculata (Walk.) Keissl. in 23, 348 (as a doubtful 
species) ; Obryzum Walk, in 56, 266; 77, 92; Verrucaria Leight. 31, 497; 
Sphaerulina Vouaux, 71 (1913), 36. On various species of Leptogium 
and recorded from five botanical counties. Description by A. L. Smith 
in 57, 289. The inclusion of this species by Saccardo in Metasphaeria 
( 1 905) and by Vouaux in Sphaerulina (1913) requires some explanation, 
as both these genera have septate spores. Apparently the spores in the 
apothecia of the host were taken instead of those of the parasite. Both 
Nylander (1858-60) and Leighton had diflSculties in regard to this, 
and the latter gave the spores as ^obsoletely 7-septate^ A. L. Smith 
( 1 9 1 1 ) described the spores as ^ simple with several guttulae ’ . Keissler, 
on account of this, puts the parasite under Guignardia. 

G« fimbriatae Keissl. 23, 346; Paralaestadia Vain. 70, i, 75 and 245. 
A specimen collected by R. Burn on Cladonia on Gader Idris. Its chief 
distinctions from other Guignardia species are its host and the negative 
coloration with iodine. 

G« insularis (Mass.) Keissl., Endocarpon Massal. 23, 343; Dermatocarpon 
insular e Migula in 76, 1933 and 1938. Recorded from six vice-counties. 
In small insulated patches on calcareous rock over Lecamra albescens 
SLiid. Placodium. For description see 76 (1933), 336. This species and 
the following (psoromoides) are given by Keissler under Guignardia^ ad 

G. psoromoides (Borr.) Keissl. 23, 346; Verrucaria Borr. 18; Endocarpon 
Leight. 28, 13; ? Laestadia Vouaux, 56, 344; Physalospora Wint. 4, 147; 
57> 385. On trunk of elm, Hurstpierpoint and Beeding, Sussex. 
Description in 42, 267, Spores given by Keissler as 12-14 x 5 ja. 

British Fungi Parasitic on Lichens. W. Watson 319 

Sphaerulina dolicliotera (NyL) Vouaux, 71; 23; 57, 386; Verrucaria 
Leight. in 31, 497; Obryzum NyL in 56, 266. Parasitic on collemaceous 
lichens, Blair Atholl. Described in 57, 289. 

S» cornicnlata Vouaux, 71 (1913), 36; Obryzum Walk. 77, 92, is trans- 
ferred by Keissler to 

S« endococcoMea (NyL) Sacc. & D. Sacc. 71 (1913), 35; 23, 436; 57, 386; 
Verrucaria NyL in 31, 493; apud Carroll, 5; Lindsay, 50 (1869), 351; 
Grombie, 8, 1 16. A. L. Smith, in 56, 344, places Verrucaria dubiella NyL 
as a probable synonym of V. endococcoidea. Keissler states that if this is 
correct, though both are described by Nylander on the same page in 
the same Flora^ dubiella precedes endococcoidea and therefore has priority, 
so that Sphaerulina dubiella is the correct designation. Parasitic on the 
thallus of Rhizocarpon excentricum, Ben Lawers, also from Killarney and 
near Dublin. For description see 31, 493. 

S. dubiella (NyL) KeissL 23, 437; Verrucaria in 31, 477, where a description 
is given; Pharcidia ? dubiella A. L. Sm. 56, 344; 4, 171. See Sphaerulina 
endococcoidea. Recorded by Carroll in 20 (1866), 25, from Ben Lawers. 
Nylander considered it to be a ‘species beni distinctaL 

Mycosphaerella cookei (Linds.) Sacc. & D. Sacc. 23, 381; Microthelia 
Linds. 39 (1869), 537, PL XXIII, f. 29. Perithecia black, scattered 
under the thicker and whitish parts of the thallus of the host [Lecanora 
crenulata). Ascus many-spored, clavate becoming irregularly swollen, 
47 X 11-5 /X, Paraphysoid hyphae ? in a slimy mass. Spores longly 
ellipsoid, hyaline, i-septate (often simple), 5-5x2*25^16. Found by 
Cooke on thallus of L. crenulata in England, 

[Astrotbelium parmularia (Berk.) Leight. 31, 499; Sphaeria Berk, in 
Hook. Journ. (1851), 19. This is given and described as a lichen by 
Leighton, with locality New Forest. A. L. Smith states that it is a 
fungus, 56, 263. It is given as Valsaria parmularia (Berk.) Sacc. in 
4, 1 75 -] 

Oiscothecium acervatum (Stirt.) A. L. Sm. 57, 387, with description; 
Lophothelium Stirt. 54 (1887), 37; 56, 265. From Ben Lawers and Killin, 
on the squamules of Stereocaulon condensatum, causing them to be 
distorted. Keissler considers that the description is incomplete. The 
paraphyses are stated to be ‘rather sparse’ in 56, but in 57 they are 
given as ‘crowded’. The latter description is probably correct. 

D. gemmiferum (Tayl.) Vouaux, 71 (1913), 46; 23, 385; 57; 44 (1935), 
235; 75, n, HI and iv; 78, 62; 77, 93; Verrucaria Tayl. in 11,11, 95, and 
28, 47 and 75; Microthelia Mudd, 42, 306; Tichothecium Krb. in 4, 175; 
56, 343; Verrucaria rugulosa Borr. 28, 47; Microthelia Mudd, 42, 306; 
Verrucaria larhalestierii Leight. 68 (1877), 242; ? V. melaspora Tayl. in 
Land. J. Bot. (1847), 153; Endococcus gemmifems Nyl. and E. rugulosus 
NyL in 8, 122. Common and widely distributed on crustaceous lichens. 
As type Keissler takes those with medium spores (8-12 x 4-6/^) ; var. 

those with small spores (6— 7 x 5-6^); var. ra/rarico/wm has 
large spores (13-22 x 6-8 ft) whilst var. physciicolum differs by the 
formation of galls. 

320 Transactions British Mycological Society 

Var. brachysponim Vouaux, 71 ; Tichothecium gemmiferum var. Zopf, etc., 
^3? 39 1* various Lecideas and other crustaceous lichens, but few 
British records. 

Var. calcaricolum (Mudd) Keissl. 23, 389; 77, 93. Discothecium calcari- 
colum Vouaux, 71 (1913), 49; Microthelia Mudd, 42, 306; Verrucariay 31, 
495; Tichothecium, 4, 175; Verrucaria perpusilla height. 31, 496; 77, 93; 
Tichothecium (Nyl.) Arn. in 4, 175; Endococcus Nyl. in 3s, 496 ; Verrucaria 
Jumosaria height. 68 (1876), 239. Endococcus .calcareus Nyl. in 8, 122. 
Has been recorded from many British localities. 

Var. physciicoium (Nyl.) Keissl. Mycoporum Nyl. 23, 392 ; Discothecium 
physciicolum Vouaux, 71 (1913)5 48. Microthelia parietinaria hinds, in 69 
(1869)3541. On Xanthoria parietina, 

D. squamarioides (Mudd) Keissl. 23, 403; 78, 62; Sphaeria Mudd, 42, i$o; 
Tichothecium Wint. in 4, 175; Sorothelia Zopf in 57, 385. Description 
of this fungus parasitic on Harriman's Teesdale specimen of Squamaria 
gelida is given in 42, 130. The perithecia are 150-250/^ diam. and the 
spores 9-25 X 5-9^6. It occurs on Phlyctis at Cleeve, v.c. 6. 

D. stereocauEcolum (hinds.) Vouaux, 71 (1913)5 57; 575 387; Micro<- 
thelia hinds. 69 (1869), 537. On the podetia of Stereocaulon in Glen 
Derrie, Braemar, v.c. 92. It is insufficiently described sec, Keissler. 
It produces deformations on the podetia forming galUike warts. 
Perithecia single or two in each wart, only emergent at apices. Ascus 
elliptic with distinct stalk, usually thick- walled, 60 x 17/i. Paraphyses 
(? periphysial hyphae) indistinct. Spores 8-nae, obovate, biseriate, 

I -septate, hyaline to olive or brown, 12-5 x 8 /i. 

D. stigma (Krb.) Zopf; Tichothecium Krb. 23, 393, 71 (1913), 52 and 
68 (1869), 367. Perithecia scattered but numerous, slightly 
bleaching or shading the host, quite sunk, ovate or top-shaped, 
slightly convex 1 20-300 ju, long by 240 /x broad, black. Ascus fusiform, 
slightly ventricose, above rounded and usually with a thick wall, 
below forming a thick foot. Periphyses present. Paraphyses (? peri- 
physoid hyphae) soon disappearing. Spores 8-nae, biseriate, i -septate, 
not or little constricted in the middle, 13-20 x 4*5-8 /x, pale olive to 
sooty brown. Hym. gel. red or blue or negative with iodine. On 
lecideoid lichens, recorded from hlanberis, v.c. 49, and Glen Dole, 
v.c. 90. 

D. vermicularium (hinds.) Vouaux, 71 (1913); Microthelia hinds, in 68 
(1869), 319 and 69 (1859)3 144 23, 407, On the thalius oiCermia 

vermicularis. Perithecia numerous, sunk, pressed together or flattened, 
very small, black (when moistened brown) with brown cells forming 
the tissue. Asci irregularly massed, obovate, unstalked, with thick 
walls at their apices. Spores longly ovate, brown, 8-nae, i-septate 
with a strong constriction, about 8-1 1 x 3-4 /X. 

Tichothecium cermariiim (Mudd) Berl. & Vogl. 4, 175;' ,'56, .344; 57, 
Sphaeria Mudd, 42, 136, is a doubtful plant which is probably 
better included in T, pygmaeum var. according to Keissler. 

Collected by Mudd from Ayton in v.c. 62, 

British Fungi Parasitic on Lichens. W. Watson 321 

T. pygmaeum Krb. Massee in 14, xvn (1888), 4, 175; A. L. Smith & Rea 
(inclusive ofvar. ventosicolum), 60; 56, 343; 57, 387; 75 (1942); 77, 93; 
78, 62; Microthelia Krb. 23, 119; M. ventosicola Mudd, 42, 307; Verru- 
caria height. 31, 495; Sphaeria ventosaria Linds. 69 (1866), 439; Endo- 
coccus ventosus Nyl. 8, 123. For description see 23, 411 and 31, 495-6. 
E. thalamitus Nyl. in Crombie, J. Linn. Soc. (1887), 217 refers to a 
foreign specimen. Keissler gives two varieties; var. ecatonosporum with 
perithecia up to 400 jw diam., asci to 95 x 23/r and spores to 12 x 6/r; 
var. erraticum with perithecia 40-90 /r, asci 30-50 ju. and spores broadly 
ellipsoid, 3-5-6 x 2-3 /r; the type being intermediate in the size of 
these organs, perithecia 50-400111, asci 45-95 x 15-23 ju, and spores 
4-12 X 2-6 /r. 

Var. ecatonosporum (Anzi) Wint., Microthelia Anzi in 23, 414. 

Var. erraticum (Massal.) Vouaux, 71; 23; 75 (1942); 77, 93; 25, 432; 
Endococcus Nyl. in 8, 122; subsp. microphons Nyl. apud Crombie in 
20 (1882), 276; Verrucaria height. 31, 496, which see for description. 
Recorded from many localities. 

Var. ventosicolum (Mudd) Wint. in 4, 175, is not considered distinct 
by Keissler. 

Pharcidia aggregata (Mudd) Vouaux, 71 (1912), 252; 23, 372; 57, 
386; 25, 341; Thelidium Mudd, 42, 289;.38 (1869), 346; 57, 386. On 
Aspicilia calcarea, Barclay’s Rock leg. Adm. Jones. On Pertusaria 
lactea on rock above Llanarmon in v.c. 50. For description see 42, 298. 

P. allogena (Nyl.) Sacc. 23, 362; 57, 386; Verrucaria Nyl. 31, 492; ArtAo- 
pyrenia Arn. 56, 324 and 57, 356 (which see for description) ; Verrucaria 
epidermidis var. apud Carroll in 5 (1886), 25. On Rhizocarpon in Ireland 
(Kylemore Castle) and on Ben Lawers. Form innata (Nyl.) Keissl. 
23, 363. Verrucaria innata Nyl. in Carroll 20 (1866), 25; 31, 494; 
Pharcidia Sacc. & D. Sacc.; P. innatula Zopf sec. 23; Vouaux in 57, 
386; Verrucaria in Hue Add. 301. For description see 31, 494. On 
Ben Lawers, parasitic on the same thallus as Pleospora kookeri. 

? P. consociata (Nyl.) A. L. Sm. Verrucaria Nyl. ex Carroll in 20 (1865), 
293. A very minute, unsatisfactory plant ‘apparently parasitic on an 
alien thallus. The spores are i-septate and broader at one end’, 57, 
386. On the summit of Ben Lawers. 

P. dispersa (Lahm) Wint. 71 (1912), 234; 23, 354; Epicymatia 

thallina (Cke.) Sacc. in 4, 176; Sphaerella Cke. 6, 372 ; Pharcidia crombii 
Sacc. & D. Sacc. in 57, 386; Endocarpon Mudd, 42, 36; Pharcidia 
frigida (Sacc.) Vouaux in 23 (also included by Keissler as it is like 
P. crombii and also occurs on Thamnolia), Pharcidia hageniae Rehm 
in 14, xvni, 79. Verrucaria conspurcans height, in i (1868), 29, with 
synonyms Arthopyrenia and Pharcidia, refers to a foreign specimen and 
is included by Keissler. For description see 42, 23, 71 and 10 (1869), 
233. Records few, Eastbourne, Talsarnau, Penmanshiel, Craig 
Rossie and near Taunton, but the fungus, which occurs on the thalli 
of many lichens, is probably frequent. 

P. dubieUa (Nyl.) A. L. Sm. 56, 344; 4, 171, is given by Keissler under 
Sphaerulina, 2-^, {s&tS. endococcoidea). 



322 Transactions British Mycological Society 

P. epicymatia (Walk.) Wint. 23, 373; 57, 386; 75> i and in; 78, 62; 
Sphaeria Walk. Pharcidia congesta Krb. as synonym in 57; 50 (1869), 
343; Epicymatia vulgaris Fkl. 4, 176; Sphaerella B. & Br. in 6; 
Sphaeria apotheciorum Massal. in 6, 872. Described in 23, 71, etc. 
Common on the apothecia of Lecanora species and recorded from many 
British vice-counties. 

P. gyrophorarnm (Arn.) Zopf, Arthopyrenia Arn.; Pharcidia gyrophorae 
(Arn.) Zopf, Arthopyrenia Arn. in 23, 379. Keissler takes the former 
name as having priority. The species occurs on Gyrophora. Sec, A. L. 
Smith in 57, 334, a minute fungus on Dermatocarpon squamules from 
Ben Lawers agrees, except in the host, with Thelidium superpositum 
{Pharcidia super posito)^ but is also akin to P. gprophorae, 

P. microspila (Krb.) Wint. in 57, 386; 75, iv; 78, 62; 25, 421-2; Artho- 
pyrenia Krb. in 56, 322; 57, 353; 75> ni; A. rhyponta Mudd, 42, 303; 
VerrucariarhypontaBoxr. (non Ach.) in 17, ii; 39, 28; 57, 353; 61, 154. 
Description in 57, 353. It occurs on or near the thallus of Graphic and 
has been recorded from five English, three Scottish and six Irish 
botanical vice-counties. Another species of Arthopyrenia^ A, spilobola 
(Nyl.) A. L. Sm. 57, 354, may also have to be included with fungi. 
Knight’s specimen from Keswick and my own from Killarney have 
very few or no algal cells. Arthopyrenia! colleta A. L. Sm. 57, 356; 
Verrucaria Stirt. 31, 468 also is probably a fungus which may be a 

P. pmctilla Wint. 23, 378; 71 (1912), 241. Keissler considers this to be 
a Pharcidia^ though it is rather insufficiently described. It may be the 
same fungus as Didymella coarctatae B. de Lesdain, 34 (1907), 695. Both 
Vouaux and Keissler agree in placing Lesdain’s species as a probable 
Pharcidia, Perithecia scattered, very small, punctiform, semi-globosely 
emergent, dark brown. Ascus from a wide-bellied base to smaller 
above, sessile with 8 irregularly ordinated spores, 30-40x14-16/^. 
Spores longly clavate, i-septate, hyaline, 14-18 x 3*5 /x with rounded 
ends and at septum not or very slightly constricted. On Biatorina 
cyrtella^ Broomfield, near Taunton, Somerset. 

P. superposita (Nyl.) Sacc. & D. Sacc. 23, 368; 71 (1912), 248; 57, 386; 
Verrucaria Nyl. in 31, 494 ; 20 ( 1 866) , 25 ; Thelidium A. L. Sm. in 5^ 300. 
Found by Carroll on Polyblastia theleodes on Ben Lawers. Described as 
Thelidium superpositum in 57, 328. 

P«? tripkractoides (Nyl.) A. L. Sm. 56, 344; 4, 171; Endococcus Nyl. ■ ex 
Crombie, 14 (1874), 24; Verrucaria Leight. 31, 497, is given by Keissler 
as synonymous with Phaespora parasitica, 

Epicymatia thallopMla (Cke.) Sacc. in 54 (1911), 38; Sphaeria Cke. in 
6, 872; Sphaerella Cke. in 14, xviir, 79; Massee, 14, xix, 44; Psilo- 
sphaeria Stevenson in 62, 388. On lichens, Scotland. The position of 
this is doubtful. It is given by Cooke in 6, 872 as ^scattered or 
gregarious, semi-immersed in the lichen thallus ; perithecia subglobose, 
carbonaceous, papillate, pierced; asci cylindrical; sporidia uniseriate, 
elliptical, uniseptate, not constricted, hyaline, at length pale greenish 
yellow, .0004 in. long. On the thallus Lecanora subfusca^ Glenshee, 

British Fungi Parasitic on Lichens. W. Watson 323 

Aug. 1856. Dr W. Lindsay. It is not improbable that this is a 
naked Sphaeria springing from the wood beneath and perforating the 
thin lichen thallus. A single small specimen was all that we have 
seen and that was insufScent to satisfy us on this point.’ Wheldon, 
in 54 (19”). 38, states ^this is evidently not the cascj as we have 
found the perithecia both on the thallus and on the apothecia of 
the lichen and from one to fourteen perithecia have been observed 
on a single apotheciumh It has been considered as synonymous with 
Pharcidia dispersa^ and the Eastbourne specimen (at one time referred 
to it) is SO5 but from Wheldon’s reference to the occurrence of the 
parasite on the apothecia of the Loch Rannock plant I am inclined 
to place his plant under Pharcidia epicymatia though I have not been 
able to examine the specimen. 

Phaeospora canmae (Ph. & Plow.) Vouaux^ 71 (1913)^ 74 i 23, 428; 
Sphaeria Ph. & Plow. 14 (1877), 27; Psilosphaeria Cke. & Plow. 14 
(1879) ? Heptameria Cke. 14, xviii, 31. Given in 4, 182, as occurring on 
Peltigera canina at Dunsley. Probably Dursley in Gloucester is the 
correct locality, as Joshua collected the material. It has also been 
recorded for Killin, v.c. 88. 

P. epicallopisma (Wedd.) FL Bad. ; Verrucaria Wedd. 23, 428. Perithecia 
semi-immersed, globose or semi-globose with gaping ostiole, quite 
black, 200-250/^ diam. Ascus probably 8-spored. Spores brown, 
3-septate, elliptical or ± ovate, 24-36 x 8-12 /l. Hymenium without 
paraphyses and not coloured with iodine. Usually occurs on Caloplaca. 
The only British record is near Taunton where it occurred on 
Placodium murorum, 

P* exoriens (Stirt.) A. L. 'Sm. 57, 388; Endococcus Stirt. 54 (1880), 220; 
Pyrenococcus Wheld. & Wils. 79, 69; 55, 482 (with description). 
A. L. Smith considers this to be worthy of segregation from Phaeospora 
parasitica^ but Keissler does not mention it. It was found at Kinloch 
Rannock, v.c. 88, on Pannaria, 

P, heteraizans Arn. 23, 427; 57, 338; Verrucaria height. 31. 493; F. mar- 
gacea in height. 28, 62 and PL XXVI, 3. Described in 31,^ 493. 
Collected by Borrer on F. submersa in Sussex. Microthelia dissepta 
A. L. Sm. 56, 352; 57, 364; 25, 426; Verrucaria Nyl. in 14 (1877), 107; 
31, 480 is now included under Phaeospora heteraizans. It was collected 
by Larbaiestier on mica-schist rocks, Doughruagh Mts., Galway, and 
by A. L, Smith on Rhizocarpon confervoides at Dooega in AchilL 
Nylander thought that the plant was probably a parasite. 

P. parasitica (honnr.) Arn. 23, 421; Verrucaria peripherica Tayl. ii, n, 97; 

Cromb. 8; Microthelia Mudd, 42, 308; Phaeospora Axny, 
Verrucaria rimosicola height. 31, 496; 69 (1869), 367; Microthelia^ 
42, Tickothecium Arn. in 56, 344; Phaeospora Zopf in 57; 78, 62 ; 
79, 93 ; 25, 402; Pharcidia triphracioides (Nyl.) A. L. Sm. 56, 344; 4, 171 ; 
Endococcus Cromb. 14 (1874), 24; Microthelia petraeicola lAxid^. sec, 
Vouaux, 71 (1913), 69; M, disjporaiorm orte/om Wats. 74 (1925)? 
Verrucccria gagei Deakin non Borr. i: (1854), 37; V. advenula Nyl. with 

324 Transactions British Mycological Society 

other references in 57, 388; 20 (1867), 260; 50 (1869), 35 ^ 1 % 12 1. 
For description see K rimosicola in 31, 496. On lecanoroid and 
lecideoid lichens, common and widely distributed. 

Var, dziednszyeMi (Bob.) Keissl. Differs from the type in having only 
two spores in the ascus. It was described as Microtkelia dispora by 
A. L. Smith in 57, 331, from a specimen collected by Joshua at 
Sapperton in Gloucestershire. A specimen collected by Knight at 
Cheltenham agreed, both externally and internally, with M. dispora^ 
except that the ascus contained 8 spores, was provisionally referred to 
fcH*m octospora in 74 (1925), 132, and given as M. dispora form octospora 
Wats, in A. L. Sm, 57, 364. Keissler considered that my plant was 
typical Fhaeospora parasitica and that Miss Smith’s was the above- 
mentioned variety. Microtkelia exerrans A. L. Sm. 56, 332; 57, 364; 79, 
72; Endococcus Nyl. apud Cromb. in 14 (1880), 1 14 and 20 (1882), 276. 
On quartzose stones at the summit of Cairn Gowar, Blair Atholl. This 
was considered by Crombie as quite possibly a fungus. So far as the 
rather incomplete description warrants, it seems to belong to Phaeo- 
spora and may be a form of the variable P. parasitica^ though the spores 
are small even for this. 

P, parmeliarum (Ph. & Plow.) Vouaux, 71 (1913), 75; 23, 430; Sphaeria 
Ph. & Plow, in 48 (1876), 124; Psilospkaeria Cke. & Plow., Melanomma 
Cke., Heptameria Cke., Leptophaeria Sacc. in 4, 1 85. On Parmelia^ Dolgelly. 

P. supersparsa Arn. in 23, 427. On the thallus of Lecidea macrocarpa^ 
Killarney. Perithecia scattered or aggregated, immersed at first, but 
later, with the blunt apex showing, broadly ellipsoid or ovoid, 
60-200 jLt diam., dark. Asci cylindrical or somewhat swollen in the 
middle, shortly stalked, 85-90 x 10- 14*5 /x. Spores 4-nae (seldom 
5-6-nae) usually with 3 septa (seldom with less or more) brown, 
± ellipsoid, often pear- or egg-shaped (seldom fusiform or clavate), 
straight or little curved, about 16-21 x 7-i2*5ft.. No definite colora- 
tion with iodine. 

P* vesicularia (Linds.) Arn., Microtkelia Linds. 69 (1869), 543 and 
PL XXIV sec. Vouaux, 71 (1913), 126. Keissler considers that this is 
a doubtful species which, on account of the many-spored ascus, 
cannot belong to Phaespora^ 23, 431. Perithecia aggregated, ± super- 
ficial, black. Ascus many-spored, (?) 68*5x21 ft. Spores 1-3-septate, 
usually 3-septate, fusiform or ovate, brown, 8-12 x 4|x. On Pertusaria^ 
Balthaycock woods, v.c. 89. 

Merismatium lopadii (Anzi) Zopf, Celidium Anzi in 23, 441. Keissler 
considers that Polyblastia nigritella Arn. (A. L. Sm. in 56, 305), Verru- 
caria Nyl. in Flora (1865), 357, Merismatium Vouaux (1913)3 77 to be 
identical. If this is so, Merismatium nigritellum (Nyl.) Vouaux seems the 
correct designation, as Nylander’s specific name dates from 1865 
whilst Anzi’s specific name of lopadii in AuL Soc. ItaL ScL dates 
from 1 868. For description, etc., see Carroll in 20 ( 1 866) ,25, Crombie, 
8, no, Leighton, 30, 466 and 31, 497 and A. L. Smith, 57, 334. The 
only British locality given is Ben Lawers, where it was with Dermato- 
carpon cinereum. A. L. Smith, in 57, 334, that Polyblastia gothica 

British Fungi Parasitic on Lichens. W. Watson 325 

Th. Fr. is similar if not identical. This was collected by Leighton at 
Shrewsbury on decaying humus and larch poles, was at first given as 
Verrucaria pituphloia height, in 30, 458 and later as V. gothica height, in 
31, 490. Th. Fries recognized that this species resembled a Sphaeria. 

[LeptorhapMs epidermidis (Ach.) Th. Fr. in 56, 330, etc. The thallus 
is very thin or often practically absent. It has been included amongst 
the fungi by Wainio {LicL Brazil^ 13) and others (73, 316), but is 
definitely placed with the lichens by most lichenologists. Our other 
species of Leptorhaphis, L, carrolii A. L. Sm. 57, 362, may be considered 
similarly. It occurs in other localities besides 'the only locality’ given 
by A. L. Smith.] 



Didymella collemata Vouaux, Cercidiospora species Stein, in 71 (1913), 
97. Keissler does not mention this in 23. On the thallus of Collema on 
rocks in Goblin Combe, Somerset, v.c. 6. SiMlar to Didymella pulposi 
but with rather larger perithecia, asci and spores, the latter 6-8-nae 
and thinning out at one or both ends, 73, 316. 

D, epipolytropa (Mudd) Berl. & Vogl. 4, 165; 23, 453; 57, 389; 78, 62; 
75 (^ 939 )> 5155 Thelidium Mudd, 42, 298; Didymosphaeria Wint. in 56, 
344; Verrucaria Cromb. 8, 12 1; 31, 494; Arthopyrenia verrucosaria 
Linds. 50 (1869), 349. On Lecanora polytropa, Aspicilia verrucosa and 
other crustaceous lichens in v.c. 19, 47, 48, 62, 88 and 96. For 
description see 31, 42, 56 and 57. 

D. pulposi (Zopf) Vouaux, 71 (1913); 78, 62; Didymosphaeria Zopf in 4, 
1745 73> 316; 77, 93. On Collema and Leptogium. As yet only recorded 
from three botanical counties, but probably much more frequent. 
Perithecia ± immersed in warts on the thallus of the host, 160-2 00 /x 
diam. Ascus clavate, about 45-75 x 10-17 ju., 4-6-spored. Spores 
longly ovate, colourless, with obtuse ends, about 13-20 x4*5-7-5/x. 

D. spMnctrinoides Berl. & Vogl. in 23, 458 and 71 (1913)5 9 ^* 
Perithecia dark brown, 150-250/x with a small ostiole. Ascus 
70-100 X io~i3*5/x. Spores 8-nae, 18-22 x 6-7*5jU.. Paraphyses 
numerous and longer than asci. On Verrucaria near Wiveliscombe, 
Somerset. Because of its host this would have been included by V ouaux 
under var. verrucariae (Zopf) Vouaux. Six varieties are given by 
Vouaux (often on account of host), but Keissler gives all of them as 
synonyms of the type. 

Metasphaeria cetraricola (Nyl.) Sacc. 4, 178 is placed by Keissler 
(23, 273), under Phragmothyrium. 

M. tartarina (NyL) KeissL, Verrucaria Nyl. in 23, 487, depends for its 
inclusion on two uncertainties, Keissler gives F. Linds, in 

68, xxvn, 343 as a probable synonym, and Lindsay’s reference 
apparently refers only to a foreign specimen. 

Didymosphaeria geUdaria (Mudd) A. L. Sm, 23, 480; 56, 344; 78, 62; 
Sphaeria Mudd, 42, 130; Tichothecium Berl. & Vogl. in Massee, 14, 
xvn; 4, 175. Collected by Harriman on the thallus of Squamaria gelida 

326 Transactions British Mycological Society 

in Teesdale and described by Mudd (spores are given by Keissler as 
mostly 4-nae and lo x 6-5^). 

D. melanospora Hepp is given by Vainio in 70, i, 15 1 as equivalent to 
Microthelia atomaria Koerb. This is given as a rare lichen by A. L. 
Smith in 56, 331 and 57, 363. It has been recorded from Cricklease 
near Chard in v.c, 5 and Kylemore in Ireland, vx. 16. 

D, microstictica Wint. 23, 475; 56, 344; 4, 174; Verrucaria height. 30, 
461; Endocarpon height. 29, no. 317, without description; Acarospora 
cervina var. microsticta Mudd, 42, 159. For description see 31, 493. On 

D. micula (Plot.) Vain, is given by Vainio in 70, i, 146 and 254 as 
equivalent to Microthelia micula (Plot.) Koerb. given by A. h. Smith 
in 56, 331 and 57, 363 as a lichen. It occurs in 20 British and Irish 
vice-counties. No definite trentepohlioid algal cells were seen in a 
Cwm Bychan (v.c. 48) specimen, but a few were noticed in a specimen 
from Ben Doran (v.c. 98). 

Leptosphaeria baeomycearia (hinds.) Sacc. & Trott, 71 (1913); Micro-- 
thelia hinds. 39 (1869), 541 or 554. On Baeomyces rufus in Britain sec. 
23, 501. Keissler considers that the description is so incomplete that it 
is better discarded, though, if it is referred to Lepiosphaeria neottizanSy the 
specific name is earlier. 

L. crozalzii (Sacc.) Vouaux, 71 (1913)5 120 and 23, 498. Verrucaria 
tartaricola hinds. 38 (1869), 351 is given by Keissler as a probable 
synonym, but the only reference found referred to a Greenland plant 
on Ochrolechia tartarea. 

L. leucomelaria (Mudd) Vouaux, 71 (1913)5 X2i and 23, 494; Sphaeria 
Mudd, 42, 105; Tichothecium &L VogL in 4, 175. On thallus of 
Anaptyckia ciliaris 3 iid A. leucomelaena. 

L* neottizans (height.) Zopf in 23, 494; Verrucaria height. 68 (1878), 
239; Didymos^aeria A. h. Sm. 56, 344; 4, 174. On thallus oi Baeo- 
mycesy Fishguard. For description see 31, 497. Vouaux has suggested 
that Microthelia baeomycearia hinds, 39 (1869), 541 or 554, might be 
included here. If so, this specific name has priority. 

L. pycnostigma (Nyl.) Sacc. & D. Sacc., Verrucaria Nyl. in 23, 495. 
Keissler considers that Microthelia baeomycearia hinds, might be refer- 
able here. 

[Massaria scoriadea Cke. in 14 (1889), 93; Sphaeria Fr. in 56, 345; 
Massariella Sacc. in 4, 174 (with other references). It was given as 
the lichen Verrucaria conferta Tayl. in ii, 185; 28, 39.] 

Pleospora ? addixbitaiis (Stirt.) A. h. Sm, 57, 328 and 389; Verrucaria Stirt. 
54 (1880), 220; Polyblastia Wheld. & Wils. 79, 71; 55, 483 (with 
description). On decaying wood, Kinloch Rannoch, vx. 88. 

P. hookeri (Borr.) Keissl. 23, 503; Verrucaria Borr., Massal, in 

56, 273; 75 (1935)5 520; Sphaeria Nyl. in 39 (1869), 548; ? Verrucaria ^ 
arctata Stirt. 65 (1879), 320; Lecidea nigropunctata Hook. FL Scot. 2nd ed. 

Menzie in herb. 75 (1935), 520; L. Schaer, apud 8, 88 and 

3^5 3t>9 ; h. sphaerica Schaer, appears to be the oldest name sec. Keissler. 
On the summit of Ben hawers. For description see 57, 296* This has 

British Fungi Parasitic on Lichens. W. Watson 327 

been considered as (i) a lichen with its own thallus, (2) an apothecium 
of a fungus parasitic on a lichen thallus. For many years it has been 
included amongst the lichens, but'Keissler and Zahlbruckner take the 
latter view. 

Neolamya peltigerae (Mont.) Theiss. & Syd., Sphaeria Mont. 23, 519. 
On thallus of FelUgera mfescens, Pont Nedd Fechan, v.c. 41 . Perithecia 
scattered, immersed, globoso-ovate, deep black, apparently without 
ostiole, ringed round by the freed layer of the cortex of the host 
thallus. Ascus clavate or longly cylindrical, to iiop, long with many 
spores which are acicular, straight or slightly bent, 50- 1 00 x 2-3 /r. with 
many oil-drops. 


Ascochytula lecanorae (Vouaux) Keissl. 23, 574. An unnamed plant 
mentioned by Lindsay in 68, xxvni, 228 as a pycnidium is considered 
as probably referable here, 23, 575. 

Dendrophoma alcicorniaria (Linds.) Vouaux in 57, 390; Microthelia 
Linds. 39 (1895), 161. On under-surfaces of squamules of Cladoma 
foliacea. Dendrobhoma podetiicola Keissl. in A. L. Sm. 67 (1910), 282, 
but Keissler (23, 551) discards this specific name as alcicorniaria has 

Diplodina Uchenodes A. L. Sm. 67 (1910), 283 is not given by Keissler. 
It was found on a lichen thallus on walnut bark at Writtle, Essex, and 
collected by Piggott. 

D. solorinaria Vouaux, 71(1914), 283; Microthelia Linds, in 38 (1869), 
349 1 57 1 390 ' Keissler puts this as probably Rhagadostoma lichenicolum 
with the Pseudosphaeriales, 23, 320. 

D. vouaxi B. de Lesd. apud 71 (1914), 288; 23, 572-3. Possibly the un- 
named parasite on the thallus of Lecanora albella given by Lindsay in 
Observations lichen. Microfungi, p. 37, and the pycnidia which Lindsay, 
in 69 (1872), 284, gives on Enterographa crassa may be placed here sec. 

Libbertella peltigerae (Lib.) Keissl. 23, 582; Zjthia Lib. apud Cke. in 
14 (1880), 83. On podetia of Cladonia, but no reference' to a British 

Lichenocomum imbricariae (AJlesch) Keissl. Coniothyrium All. in 23, 
564; ? Microthelia cargiliana Linds, in 69 (1866), 439 and Pi. XXX, but 
Lindsay gives no British locality. 

Pboma abietinae Vouaux, 71 (1914), 547. On the thallus ot Lecanactis 
abietina. This depends on a very incomplete description by Lindsay 
and is a very doubtful species (23, 589). ^ ^ ^ ^ ^ ^ 

P, lecanorae Vouaux, 71 (1914), 547 j 277; 33, 164; 23, 544. On 
■ apothecia of Lecanora subfusca (agg.), Tatmton, v.c. 5, Blair Atholl, 
v.c. 89. It is probably frequent. According to Vouaux some of 
Lindsay’s spermogonia may belong to it. ‘Perithecia’ dark, + im- 
mersed, minute, 80— 150 /X diam., somewhat lens-shaped. Conidia 
about 3-7 X f5-2-5ju, ellipsoid, hyahne, borne on longer carriers. 

328 Transactions British Mycological Society 

P. uncialicola (Zopf) Vouaux, yi (1914), 1^%', Phyllosticta Zopf in Sacc. 
Syll. fung. I (1906), 245. Abrothallus moorei Linds, was included by 
Vouaux under this, but Keissler points out that Lindsay gave an 
ascus with spores to his species. See A. moorei on p. 312. 

Vouauxiella lichenicola (Linds.) Petr. & Syd. 23, 565 ; Torula Linds. 
39 (1868-9), 1 15 and 530 and PL XXIII; Sirothecium Keissl. in 71 
(1914); 15, n, 133; 72, 82. Pycnidia loosely scattered, ovate, 
ellipsoid or almost round, mostly about 60-90/4, their covers con- 
sisting almost entirely of the darkened and somewhat diminished cells 
of the host; conidiophore dark; conidia toruloid. On the thallus and 
apothecia of many lichens, chiefly Lecanoras. Apparently common, 
as it is recorded from many localities in v.c. 5, ii, 17,64, 72,81,88, 92, 
93, 98, 100; Ireland, 1-5. 


Lichenophoma Keissl. Vouaux in 71 (1914) considered that a lichen 
parasite on Biatorina {Lecidea) griffithii described by Lindsay in 68 
{iQ'js), 263, belonged to this genus, but Keissler is doubtful, 23, 589. 


Aegerita physciae Vouaux, 71 (1914), 313. Mycelium web-like, effuse, 
whitish in colour, but becoming darker and finally destroying the 
tissue of the Pfyscia. Hyphae hyaline, 3-7/4 thick, in some places 
forming a rose-tinted cushion (sporodochium) . Conidiophores simple, 
long, 25-35x4-7/1. Conidia terminal, io-15/t diam. Keissler 
(23, 628) suggests that it may be a state of Corticium centrifugum. 
No definite British records, but owing to its external resemblance to 
Illosporium roseum may have been included in the records of that species. 

Atractium flammeum Berk. & Rav. 3, 461; 41, in, 452. Keissler states 
that it breaks through the underside of Parmelia subaurifera, Physcia 
stellaris and Xanthoria and is considered to be the conidial stage of 
Sphaerostilbe flammea Tul. 46, 262. It is given as Microcera coccophila 
Desm. in 72, 67. Described in 41. 

Coniosporium physciae (Kalchb.) Sacc., Gymnosporium Kalchb. in 23, 
604; 41, ni, 357; 72, 71. Given by Phil. & Plow, in 14, iv (1876), 1 19, 
as found on Physcia at King’s Lynn, Norfolk. Its common host is 
Xanthoria parietina, but it also occurs on Squamaria, Physcia and Ramalina. 
When on the last-named host it is given as Spilomium ramalinae Oliv. in 

71 (1914)- 

Comothedum ' grapMdeorum (NyL) Keissl. 233 .. Spilomium Mjl, ia 

57^ 267; 77, 93 {Spiloma melaleucum Ach., S» versimlor Smith., S. vario*^ 
losum Turn. & Borr., S, fuliginosa Turn., Coniothecium nigrum Lam., 
C, olivaceum DC. are prior names before the starting-point of fungal 
nomenclature). It forms somewhat longish and irregular groups on 

British Fungi Parasitic on Lichens. IV. Watson 329 

the host which it decomposes and blackens. The conidia are ± muri- 
form, irregularly globose, 6-8 ji* diam. On Opegrapha atra and O. lyncea 
and noted from v.c. 5, 6, 17, 56. 

C. Uchenicolum Linds. 39 (1869), 518 and 534; Spilomium Vouaux, 71 
(1914), 321. OntYizShisoi Bmllia,Lecanoraz.-a. 6 iPertiisaria, and recorded 
from various localities in v.c. 38, 40, 48, 64, 78, 83, 85, 88, 89, 92, 104, 
107; Ireland, i, 2, 38. Forms wart-like bodies, punctiform and black. 
The conidia are 6-12 x 6/i, dark brown, at first ± globose and later 
+ conical. 

C. sUaceum (Linds.) Keissl. 23, 619; Gassicurtia Fee in Linds. 39 (1869), 
542; Spilomium Nyl. in Vouaux, 71 (1914), 322. Similar to Coniothecium 
graphideorum but not on Opegrapha, and the conidia contain oil-drops. 

C. sphaerale (Fr.) Keissl. 23, 616; 75, iv, 242; 78, 62; Sclerococcum Fr. in 
67 (1917)5 433; Acolium corallimm Krb. in 38 (1869), 342. On the 
thallus of various lichens, especially species of Pertusaria. Some 
specimens attributed to Cyphelium stigonellum A. Zahl. and C. notarsii 
A. Zahl. in 55, 22, and in other British lichenological works, belong 
here. Pertusaria corallina, when attacked by this parasite is ‘sometimes 
referred to a distinct form, f. papillosa (Ach.) Zahlbr.’, 27, 318. The 
type is probably common in the British Isles and has been recorded 
from many botanical vice-counties. 

Dactylium lichenicolum Vouaux, 71 (1914), 307, is probably a 

saprophyte as it occurred on the decayed thallus of a Parmelia. It is 
placed by Karst, as a subspecies of Dactylium dendroides Fr. which is 
given in 41, nr, 341, without any definite reference to its occurrence 
on lichens. 

[Epicoccum neglectom Desm. in 41, nr, 488 ; 72, 99 ; occasionally occurs 
as a saprophyte on Peltigera and other lichens.] 

Fusarium kuhxdi (Fkl.) Sacc. in 41, m, 484; 72, 65; Fusisporium Fkl. in 
14 (1876), 1 20, where it is given as occurring on mosses and lichens, 
Norfolk. Keissler’s only reference to this species is as a state of 
Coriicium centrifugum, 

Hjphoderma roseum Fr. 72, 74 with other references; Hyphelia Fr. It is 
considered by Keissler to belong to Corticium centrifugum. 

IllosporiiiBi carneum Fr. 41, in, 468; 72, 57. On species o? Peltigera. 

L coccineam Fr. in 41, m, 468; 72, 57. On Endocarpon, Pertusaria, 
Lecanora, etc. 

L rosenm Mart, in 41, ni, 468; 72, 57; 77, 93. On Physcia, Anaptychia, 
Parmelia, Xanthoria and Solorina. 

Var. corallinnin (Rob.) Ferr. 23, 632 ; 72, 57. On similar lichens as the type. 

[Macrosporinin commime Rabh. in 41, in, ^.^lyAlternaria tenuis ex 
Wallr. in 72, 98 with other references. This sometimes occurs as a 
on Ocholechia par ella.'l 

Spegazxinia Sacc, in Michelia, n (1880), 37. A species of this genus was 
reported as parasitic on Pertusaria at Minehead, v.c. 5. Report of the 
Brit. Myc. Soc. meeting at Minehead, 1920. This probably Conio- 
thecium sphaerale which is found on Pertusaria in the Minehead district. 

330 Transactiom British Mycological Society 

Sporotricliiim Mchejiicoliim Berk. & Br. in 21 (1872)5 1025 is given by 
Keissler as a state of Corticum centrifugum. No British locality given. 
[Trichothedum rosenm Link, in 23, 596; Dactjlium Berk.; 4I5 iii, 337; 
72, 875 with other references. Sometimes occurs on Stereocaulon ziA 
other lichens, but is a common mould and not a lichen parasite.] 


The following names are difficult or impossible to arrange in a systematic 


Caliduni arenariuin Nyl. in 55, 12; C. ciirimm Leight. in 31, 44, is 
parasitic on the thallus of Biatora lucida (Ach.) Fr., but is included by 
most authors amongst the lichens. 

Cliostomum corrugatum Fr. in 28, 69 is given as a parasite on the crust 
oiLecideaehrhariianaAdi. Biatorina graniformis A. L. Sm.). According 

to A. L. Smith (57, 129) the apparent fructifications are the spermo- 
gones of the host. 

Leddea imponens Leight. in 68 (1876), 238; 31, 385; 56, 104, which is 
parasitic on the thallus of Lecanora polytropa at Fort Flill, Fishguard, is 
possibly Ji'esolechia vitellinaria Rehm, but is given as a lichen in 57, 1 1 1 
and other places cited above. 

‘Opegrapha epiphega’. Lichen vugaszcs, Eng. Bot. On trees near the River 
Noran, v.c. 90 leg. Don and given on Turner’s authority. The specimen 
in the herb. Edinburgh so labelled is the fungus, Dichaena faginea Fr., 
for which the above are old names, as it was confused with lichens, 
755 h 519- Other Hysteriales, especially Hysterium angustatum A. & S. 
and H. pulicare Pers., are often mistaken for species of Opegmpha or 
other Graphidiales. Gloniopsis levantica Rehm, when associated with 
a lichen thallus, is liable to be mistaken for a Dictyographa. On the 
other hand, mistakes may be made in an inverse direction. For 
example, Gloniopsis watsoni Rilstone, which was described in the 
J. of Botany (1940), 192 as occurring on an indeterminate lichen 
thallus is a form of the lichen Graphina MuII-Arg., with most 

apothecia longer than usual and with the algal cells of the thallus 
more obscurely trentepohlioid. 

Pyrenothea aphanes Leight.; Verrucaria Borr. in 28, 67; 56, 202; 57, 

P. leucocephala Fr. in 28, 65; Sphaeria Pers. in 56, 202; 57, 223. 

P« litMna Leight., Verrucaria TayL in 28, 68; 56, 296. 

P. lutea Leight. in 28, 68; 56, 296. 

P. moUis Leight., Verrucaria TayL in 28, 67; 56, 296. 

P* niveoatra Leight., Borr. in 28, 67 ; 56, 296, 

P.mdis Leight., Verrucaria Borr. in 28, 66; 56, 202; 57, 223. 

P. stictica Fr. in 56, 205; 57, 226. 

P. sulphurea Leight. in 28, 69; 56, 296. 

P, vermicellifera Leight... in 29, no. 292; 56, 202;„57, 223. „ . 

British Fungi Parasitic on Lichens. W. Watson 331 

All the above-mentioned Pyrenothea {Sphaeria and Vermcaria) are 

considered to be spermogonia of Lecanactis ahietina^ Platygrapha periclea^ 
0 pegrapha and other lichens, 

TtromMuin epigaeum Wallr,, Sphaeria Pers. and other synonymy in 57, 
3365 is considered as a true lichen. Acharius’s subgenus Inoderma was 
raised to generic rank by S. F. Gray, who included in it two British 
species, I, epigaea and L byssacea^ the latter of doubtful position based 
on Sphaeria byssacea Weig. and considered by some authors as a 
spermogonial form, 57, 335. 

Verrucaria conturmatiiia Nyl. in 14, viii, 29; V, elachistophora Nyl. in 
31, 454; ¥. harrimatmi Ach. in 28, 63; V. pulposa Leight. in 30, 
427; 3I5 457, are given by A. L. Smith as parasitic or doubtful 
species, 56, 295-6; 57, 321-2. 


1 Annals and Magazine of Natural History. 

2 Berkeley, M. J. & Broome, G. E. In J. Linn. Soc. (1872), etc. 

3 Berkeley & Ravenni. In Ann. Mag. nat. Hist. (1859), 461. 

4 Bisby, G. R. & Mason, E. W. List of Pyrenomycetes recorded for Britain. Trans. Brit. 

myc. Soc. xxiv, pt. ir (1940), 

5 Carroll, Isaac, or Jones, T. In Nat. Hist, and Quart. J. Sci.f DubliU} 1859 and other 


6 Cooke, M. G. Handbook of British Fungi (1871). In Grevillea^ 1885, etc. 

7 Cooke & Plowright. In Grevillea (1879), 85. 

8 Crombie, J. M. Lickenes Britannica (iSjo). 

9 Crombie, J. M. British Lichens^ pt. i (1894). 

10 Crombie, J. M. Notes in J. Bot., Lond. 

11 Flora Hibernica. 

12 Grelet, Abbe L. j. In Bull. Soc. mycol. Fr. (1926), 207. 

13 Greville, R. K. Scottish Cryptogamic Flora. 

14 Grevillea. 

15 Grove, W. B. British Stem- and Leaffungi. 

16 Grove, W. B, Notes in J. Bat., Lond. (1912), 91, etc. 

17 Hooker, W. J. British Flora. 

18 Hooker, W. J. English Flora. 

19 Hooker, W. J. Flora Scotica. 

20 Journal of Botany. 

21 Journal of Linnea?i Society {Bot). 

22 Journal of Ecology. 

23 Keissler, K. VON. JDie Flechienparasiten (igso). Rdihcnhorsty Krypt. -Flora. 

24 Knight, H. H. Notes in Trans. Brit. myc. Soc. 

25 KInowles, M. G. The Lichens of Ireland. Proc. Roy. Irish Acad. (1929). 

26 Lamb, I, M. Lichenological Notes. J. Bot.^ Lond., 1936-41. 

27 Lamb, I, M. A lichenological excursion to the west of Scotland. Trans, bot Soc. Edin. 

■ xxiii,,.pt. in (1942). 

28 Leighton, W. A. Angiocarpous Lichens (1851). 

29 Leighton, W. A. Lichenes Brit, exsicc. 

30 Leighton, W. A. ist. ed. (1870); 31, 3rd ed. (1879). Most references are 

tothe third 'edition., ■' 

32 Lesdain, B. de. Recherches sur les Lichens des Environs de Dunkerque (1910). 

33 Lesdain, B. de. Recherches sur les Lichens des Environs de Dunkerque (1910). 

Supplement ( 1 9 1 4) . 

34 Lesdain, B. DE. Notes iichenoiogiques. Bull. Soc. hot. Fr. 

35 Lindsay, L. Monograph of the genus J. Micr. Sci. y 


Transactions British Mycological Society 

36 Lindsay, L. Memoir on the Spermogones and Pycnides of Filamentose, Fruticulose 

and Foiiaceous Lichens. Tram. Roy. Soc. Edinb. 

37 Lindsay, L. Notes in Trans. Linn. Soc. Land. (Bot.) 

38 Lindsay, L. Notes in Quart. J. Micr. Set. ^ 

39 Lindsay, L. Notes in Trans. Roy. Soc. Edinb. 

40 Mason, E, W. See Bisby. 

41 Masses, G. British Fungus Flora 

42 Mudd, W. Manual British Lichens (iS 6 i). 

43 Mudd, W. British Cladoniae (1865). 

44 Naturalist, The. 

44^ Olivier, H. Les principaux parasites de nos Lichens frangais. Bull. Acad. int. Geogr, 
bot. (1905-7)* 

45 Fetch, T. Notes, on British Hypocreales. J. Bot. (1936), 185-6. 

4 ^ Fetch, T. British Hypocreales. Tram. Brit. myc. Soc. (1938). 

47 Phillips, W. A Manual of the British Discomycetes (1887). 

48 Plowright, G. B, Notes in Grevillea (1876), 126, etc. 

49 Proceedings of the Somersetshire Arch, and Nat. Hist Soc. 

50 Quarterly Journal of Microscopical Science. 

51 Ramsbottom, j. a List of the British species of Discomycetes. Tram. Brit. myc. Soc. 


52 Rea, Carleton. British Basidiomycetes (ig 22 ). 

53 Sampson, K, List of British Ustilaginales. Trans. Brit. myc. Soc. xxiv, pts. in and tv ( 1 940) . 

54 Scottish Naturalist. 

54 Smith, A. Lorrain. Trans. Brit. myc. Soc. m (1910). 

55 Smith, A. Lorrain. British Lichem, pt. i, 2nd ed, (1918). 

56 Smith, A. Lorrain. British Lichens, pt. 2, ist ed. (1911). 

57 Smith, A. Lorrain. British Lichem, pt. 2, 2nd ed. (1926). 

58 Smith, A. Lorrain. Lichem, Cambridge Botanical Handbook (1921). 

59 Smith, A. Lorrain. Notes in Tram. Brit. myc. Soc. (1917)5 48 (with Ramsbottom). 

60 Smith, A. Lorrain. Notes in Tram. Brit. myc. Soc. ii (1899), 61 (with Rea). 

61 Smith, J. E. English Flora. 

62 Stevenson, J, Mycologia Scotica. 

63 Stirton, j. Notes in Scottish Naturalist. 

64 Stirton, J. Notes in Grm&a. 

65 Stirton, J. Notes in Proc. Phil. Soc. Glasg. (1879), 320. 

66 Tramactiom of the Botanical Society of Edinburgh. 

67 Tramactiom of the British Mycological Society. 

68 Tramactiom of the Linnean Society. 

69 Tramactiom of the Royal Society of Edinburgh."^ 

70 V41NIO, E. A. (formerly Wainio). Lichenographia Fennica, vois. i-iv, the last volume 

edited by Bernt Lynge. 

71 VouAux, Abbe. Synopsis des champignons parasites de Lichens. BmIL Soc. mycoL Fr. 


72 Wakefield, E. M. & Bisby, G, R, List of Hyphomycetes recorded for Britain. Tram. 
Brit. myc. Soc. xxv, pt. i (1941). 

73 Watson, W. New, rare and critical lichens. J. Bot., Lond. {igij). 

74 Watson, W. Lichenoiogical Notes, i-x. J. Bot., Lond. {ig2^-42). 

75 Watson, W. Notes on Lichens. Tram. Bot, Soc. Edinb. i-iv (1935-95 1942, 1945). 

76 Watson, W. Botanical Notes. Proc. Somersetsh. arckaeol. nat. Hist. Soc. 

77 Watson, W. The Lichens of Somerset. Proc. Somersetsh. arckaeol. nat. Hist. Soc. ( 1 928-30) . 

78 Watson, W. The Lichens of Yorkshire- Naturalist (1946). 

79 Wheldon, J. A. & Wilson, A. The Lichens of Perthshire. J. Bot., Lond., SiippL (1915)* 

80 Wheldon, J. A. & Wilson, A. The Flora of West Lancashire (1903). 

81 A. The Flora of Westmorland. See also Wheldon. 

82 Withering, W. Botanical Arrangement. 

83 Zahlbrugkner, A. Catalogue Lichenes Universalis. 

* Some mistakes may occur in the references No. 68 and 69. Some of the copies were im- 
perfectly “spined ” so that mistakes in number of volume or year of issue were easily made. 

British Fungi Parasitic on Lichens. W. Watson 333 


Most of the lichen-hosts are omitted from this list 

abietina Krb. (Lecanactis), 331 
abietinae (Pers.) Zahi. (Xylographa), 315 
Vouaux (Phoma), 327 
Abrothallus DN.j 309, 312, 328 
abrothallus Linds. (Gonida), 308 
Agarospora MassaL, 317, 326 
acervatum A. L. Sm. (Discothecium), 319 
Stirt. (Lophothelium), 319 
Agolium Ach., 329 

addubitans A. L. Sm. (? Pleospora), 326 
Stirt. (Verrucaria), 326 
Wheld. & Wils. (Polyblastia), 326 
Adelogogcus Theiss. & Syd., 317 
adjuncta Th. Fr. (Bueilia), 31 1 
Ax'n. (Karschia), 31 1 
advenula A. L. Sm. (Bueilia), 31 1, 312 
Leiglit. (Lecidea), 31 1 
Zopf. (Karschia) ,311,312 
Aegerita Pers. Fr., 328 
affinis Cke. (Nectria), 317 
Grev. (Sphaeria), 317 
(Grev.) Sacc. (Paranectria), 319 
aggi'egata Vouaux (Pharcidia), 321 
aggregatum Mudd (Theiidium), 321 
Agyrium Fr., 307, 315^ 
alcicornaria Linds. (Microthelia), 327 
Vouaux (Dendrophoma), 327 
aliogena Arn. ( Arthopyrenia) , 321 
Carroll (Verrucaria var.), 321 
.NyL (Verrucaria), 321 
Sacc. & D. Sacc. (Pharcidia), 321 
alpestris Theiss, & Syd. (Addococcus), 317 
Alternaria Nees ex Walk., 329 
angustatum A. & S. (Hysterium), 330 
Antennaria Lk., 307 
aphanes Borr. (Verrucaria), 330 
Leight, (Pyrenothea), 330 
aporea Nyi. (Psora), 316 
apotheciorum MassaL (Sphaeria), 322 
arachnoideum Berk, (Gorticium), 307 
Lev. (Rhizoctonia), 307 
arctata Stirt. (Verrucaria), 326 
arenarium NyL (Galicium), 330 
arenicola A. L. Sm. (Bacidia var.), 31 1 
Leight. (Lecidea), 311 
Mudd (Raphiospora) ,311 
Sacc. (Mycobacidia), 31 1 
Arthonia Ach., 307, 308, 310, 312, 317 
Arthopyrenia Mass., 321, 322, 325 
Asgochytuea Pot. emend. Died., 327 
associata Th, Fr. (Lecidea), 308 
Sacc. & D. Sacc, (Nesolechia), 308 
Astrothelium Eschw,, 319 
ater Linds. (Abrodiallus var.), 312 
atomaria Krb, (Microthelia), 326 
athallina A. L. Sm. (Bueilia form), 312 
Mull. (Bueilia), 312 
Vouaux' (Karschia), 312 
Atragtium Link., 328^ . 
atricola Linds, (Microthelia), 317 
Sacc. (Muliereila), 317 

aurantiaca B. & C. (Stigmatella), 306 
aurantiacum Kalchb. (Polycephalum), 306 
aurantiacus Thaxt. (Ghondromyces), 306 

babingtonii Berk. (Strigula), 316 
Bacidia DN., 310, 317 
Badhamia Berk., 306 
baeomycearia Linds. (Microthelia), 326 
Sacc. (Leptosphaeria), 326 
Bertia DN., 318 
Biatorella DN., 314 
Biatorina Mass., 310, 322 
Bilimbia DN., 31 1 

brachysporum Vouaux (Discothecium var.), 

319* 320 

Zopf (Tichothecium var.), 319, 320 
Broomella Sacc., 316 
bryophila Wats. (Stenocybe), 314 
bryophilus Zahl. (Diploschistes), 315 
Buellia DN., 31 1, 312 
byssaceaWeig. (Sphaeria), 331 
S. F. Gray (Inoderma), 331 

calcareus NyL (Endococcus), 320 
calcariccolum Arn. (Tichothecium), 320 
Keissl. (Discothecium var.), 319, 320 
Vouaux (Discothecium), 320 
calcaricola Leight. (Verrucaria), 320 
Mudd (Microthelia), 320 
Galicium Pers., 312, 313 
Galloria Phil., 308 
Calonectria DN., 316 
campestris Th. Fr. (Biatorella), 314 
campsteriana Linds. (Verrucaria), 325 
caninae Gke. (Hepameria), 323 
Cke. & Plow. (Psilosphaeria), 323 
Plow. & Phil. (Sphaeria), 323 
Vouaux (Phaeospora), 323 
capsulifera Berk. (Badhamia), 306 
cargiliana Linds. (Microthelia), 327 
carneum Fr. (Illosporium), 329 
carrolii A. L. Sm. (Leptorhaphis), 325 
Catillaria (Mass.) Th. Fr., 310 
Celidiopsis Sacc., 308 
Celidium TuL, 307, 308, 324 
centrifuga MassaL (Opegrapha), 313 
Stein, or Rehm. (Leciographa), 313 
centrifugum Bresad (Gorticium), 307, 329, 330 
Lev. (Rhizoctonia), 307 
Gercidiospora Korber, 316, 325 
cerinaria Mudd (Sphaeria), 320 
cerinarium Berl. & Vogl. (Tichothecium), 

cetrariicola Arn. (Nesolechia), 309 
Cke. (Psilosphaeria), 316 
Linds. (Lecidea), 309 
NyL (Sphaeria), 316 
Sacc. (Metasphaeria), 316, 325 
cetrariicolum Keissl. (Phragmothyrium), 316 
Ghaenothega Th. Fr., 312 
Ghondromyces Berk. & Curt, 306 

Transactions British Mycological Society 


chrysantheus Am. (Abrothallus form), 313 
Stein. (Abrothallus), 313 
CiLioMYCES Hohn., 317 
citrineila Ach. (Lecidea), 311 
citrinum Leight. (Galidum), 330 
dadoniaria Arn. (Nesolechia), 309, 312 
Nyl. (Lecidea), 309 
cladoniarum Linds, (Abrothaiius), 309 
ciemens Mass. (Gonida), 308 
Till. (Phacopsis), 308 
Gliostomum Fr., 330 
coarctatae B. de Lesd. (Didymella), 322 
Keissl. (? PHarcidia), 322 
coccinella Fr. (Galloria), 308 
liarst. (Orbilia), 308 
Somm. (Peziza), 308^ 
coccineum Fr. (IlJosporium), 329 
coccophiia Desm. (Microcera), 328 
Gollema Wigg., 318 ^ 
collemata Vouaux (Didymella), 325 
colleta A. L. Sm. (Arthopyrenia?), 322 
Stirt. (Verrucaria), 322 
commune Rabh. (Macrosporium), 329 
conferta Tayl. (Verrucaria), 326 
congesta Krb. (Pharcidia), 322 
CoNiANGiuM Fr., 315 
Gonida MassaL, 307, 308, 317 
CoNiosPORiUM Link ex Fr., 328 
Goniothegium Gorda, 328, 329 
Goniothyrium Gorda, 327 
cookei Linds. ( Micro thelia), 319 
Sace. (Mycosphaerelia), 319 
CoRTicruM Fr., 307, 329 
corticola Nyl. (Schizoxylon), 315 
Fr. (Goniangium), 315 
consociata A. L. Sm. (Pharcidia), 321 
Nyl. (Verrucaria) ,321 
conspurcans Leight. (Verrucaria), 321 
(Arthopyrenia), 321 
conturmatula Nyl. (Verrucaria), 331 
corallinum Krb. (Acolium), 329 
Ferr. (Illosporium var.), 329 
Rob. (Illosporium), 329 
corniculata Keissl, (Guignardia), 318 
Leight. (Vermcaria), 318 
Sacc. (Metasphaeria), 318 
Vouaux (Sphaerulina), 318, 319 
Walk. (Gbryzum), 318 
corrugatum Fr. (Cliostordum), 330 
Graterium Trentip., 306 
cristata A. L. Sm. (Biatorina), 310 
Arn. ( Gatillaria) ,310 
Leight. (Lecidea), 310 
Sacc. & D. Sacc. (Scutula), 310 
crombii Mudd (Endocarpon), 321 
Sacc, & D. Sacc. (Pharcidia), 321 
' crozalzii Vouaux (Leptosphaeria), 326 
Gyphelium Ach., 329 

'Dacampia ^Mass., , 326 ' 

Dactyeium Nees., 329, 330 
DactylospOra Krb., 313 
dendroides Fr. (Dactylium), 329 
Dendrophoma Sacc., 327 
Dermatogarpon Eschw., 318 

destructans Tobl. (Karschia), 312 
Dialonegtria Gke., 316 
Dighaena Fr., 330 
Dictyographa Miill-Arg., 330 
Didymella Sacc., 325 
Didymium Schrad., 306 
Didymosphaeria Zopf, 325, 326 
difformis Wain. (Biatoreiia), 314 
Diplodina Westend, 318, 329 
Diploschistes Norm., 315 
DiscocEiiA A. L. Sm. & Ramsb., 309 
Disgothecium Zopf, 319, 320 
dispersa Wint. (Pharcidia), 321, 323 
dispora A. L, Sm. (Microthelia), 323 
dissepta A. L. Sm. (Microthelia), 323 
Nyl. (VeiTucaria), 323, 324 
dolichotera Leight. (Verrucaria), 319 
Vouaux (Sphaerulina), 319 
Dothidea Fr., 317 
dubia Gke. (Homostegia) , 307 
Linds. (Ph>Tnatopsis), 312 
dubiella A. L. Sm. (? Pharcidia), 319, 321 
Keissl. (Sphaerulina), 319, 321 
Nyl. (Verrucaria), 319 
dubium Linds. (Celidium), 307 
duriae M. & B. (Myriangium), 318 
dzieduszyeki Keissl. (Phaeospora var.), 324 

ecatonospora Anzi. (Microthelia), 321 
ecatonosporum Wint. (Tichothecium var.), 

ecrustacea A. L. Sm. (Diploschistes form), 315 
Gromb. (Lecanora form), 315 
effusa Auersw. (Bilimbia), 31 1 
ehrhartiana Ach. (Lecidea), 330 
elachistophora Nyl. (Verrucaria), 331 
endocarpicola Keissl. (? Celidium), 310 
Linds. (Lecidea) 310 
Vouaux (Mycobilimbia), 310 
Endocarpon Hedw., 318, 321, 326 
endococcoidea Nyl. (Verrucaria), 319 
Sacc. (Sphaerulina), 319 
Endogoccus Nyl., 317, 319, 321, 323 
epibiastematica A. L, Sm. (Biatorina), 310 
Rehm. (Scutula), 310 
Walk, (Peziza), 310 
epicallopisma Am. (Cercidiospora), 316 
FL Bad. (Phaeospora), 323 
Sacc. (Nectria), 316 
Wedd. (Verrucaria), 323 
epicladonia Arn, (Biatorina), 310 
Nyl. (Lecidea), 310 
Sacc. (Scutula), 310 
Vouaux (Gonida), 310 
Epigogcum Link., 329 . 

Epicymatia Pki., 322 ■ . , ■ 

epicymatia Walk. (Sphaeria), 322 ■ 

Wint. (Pharcidia), 322, 323 
epidermidis Tli. Fr . ■ ( Leptorhaphis) , ■ ■ 325' : 
epigaea S. F. Gray (Inoderma), 3.31 ' ■ 
epigaeum Wallr. (Thrombium),"33i, 
epipasta Stirt. ■■ (Lecidea), 307 
epiphega Eng* Bot. (Opegrapha), 330 ■ 
epiphorbia Stirt. (Lecidea),' 308 . 

Vouaux (Gonida), 308" 

Zopf (Karschia), 308 


British Fungi Parasitic on Lichens. W. Watson 

epipolytropa Berl. & Vogl. (Didymella), 325 
Gromb. (Verrucaria), 325 
Wint. (Didymosphaeria), 325 
epipolytropum Mudd (Thelidium), 325 
epipsila Nyl. (Lecidea), 312 
Oliv. (Buellia)j 312 
episema A. L. Sm. (Biatorina), 310 
Nyl. (Lecidea), 310 
Oliv. (Catillaria), 310 
Zopf (Scutula), 310 
epithailina Boud. (Mollisielia), 308 
Phil. (Mollisia), 308 
Phil. & Plow. (Peziza), 308 
Sacc. (Pezizeila), 308 
epithallinum Leight. (Theiocarpon), 317 
erratica Leight. (Verrucaria), 321 
erraticum Mass. (Tichothecium), 321 
Vouaux (Tichothecium var.), 320, 321 
erraticus Nyl. (Endococcus), 321 
Eurotium Link ex Fr., 315 
euspora Anzi (Stenocybe), 314 
exerrans A. L. Sm. (hlicrothelia), 324 
Nyl. (Endococcus), 324 
exoriens A. L. Sm. (Phaeospora), 323 
Stirt. (Endococcus), 323 
Wheld. & Wils. (Pyrenococcus), 323 

faginea Fr. (Dichaena), 330 
ferdinandezii Gromb. (Lecidea), 312 
fimbriatae Keissl. (Guignardia), 318 
Vain. (Paralaestadia), 318 
firmatum Nyl. (Odontotrema), 315 
flammea Tul. (Sphaerostilbe), 328 
fiammeum Berk. & Rav. (Atractium), 328 
flavovirescens Anzi (Bacidia), 311 
Borr. (Lecidea), 31 1 
Dicks. (Lichen), 311 
Krb. (Raphiospora), 31 1 
Rehm (Mycobacidia), 31 1 
fossarum Rehm. ex Ramsb. (Biatorella), 314 
Th, Fr. ex A. L. Sm., 314 
frigida Vouaux (Pharcidia), 321 
fuistingii Krb. (Lahniia), 31 1 
fuHginosa Turn. (Spiioma), 328 
fuivus Jahn. (Myxococcus), 306 
fumosaria Leight. (Verrucaria), 320 
Fusarium Link., 329 

fuscopurpurea Vouaux (Gonida), 308, 317 
fuscopurpureum TuL (Celidium), 308 
Fusisporium auct. ant., 329 

gagei Deak. (Verrucaria), 323 
galactinaria Am. (Gonida), 308 
Leight. (Arthonia), 307, 308 
Vouaux (Gonida), 309 
Gassicurtia F^e, 329 
gelasinata Zahl. (Sphinctrina) ,314 
geiasinatus Wahl. (Lichen), 314 
gelida (Squamaria), 325 
gelidaria A. L. Sm. (Didymosphaeria), 325 
Mudd , (Sphaeria), -325 '' 
geiidarium B. & G. (Tichothecium), 325 
gemmifera Mudd (Microthelia), 319 
Tayi. (Verrucaria),. 319 
gemmiferum Krb. (Tichothecium) ,319 
Vouaux (Discothecium), 319 

gemmiferus Nyl. (Endococcus), 319 
geophana Fr. (Biatora), 309 
Nyl. (Lecidea), 309 " 

glaucomaria A. L. Sm. (Leciographa), 307 
Nyl. (Arthonia), 307 
Nyl. 1852 (Lecidea), 307 
glebulosus Sm. (Lichen), 310 
glomerulosum Tayl. (Goliema), 318 
Glonxopsis, 330 

gothica Leight. (Verrucaria), 325 
Th. Fr. (Polyblastia) , 324 
graphideorum Keissl. (Comothecium), 328, 329 
Nyl. (Spilomium), 328 
Guignardia Viala & Rav., 318, 319 
Gymnosporium ? Kalchb., 328 
gyrophorae Arn. (Arthopyrenia), 322 
Zopf (Pharcidia), 322 
gyrophorarum Arn. (Arthopyrenia), 322 
Zopf (Pharcidia), 322 

hageniae Rehm (Pharcidia), 321 
haplotella Am. (Mullerella), 317 
. Leight. (Vermcaria), 317 
haplotellus Nyl. (Endococcus), 317 
harrimanni Ach. (Verrucaria), 331 
heerii Hepp (Lecidea), 310 
Heptameria Rehm. & Thuem., 323 
herbariorum Link. (Eurotium), 315 
heteraizans Am. (Phaeospora), 323 
Leight. (Verrucaria) 323 
Homostegia Fki., 307 
homostegia Linds. (Dothidea), 317 
hookeri Borr. (Verrucaria), 326 
Keissl. (Pleospora), 321 
Linds. (Sphaeria), 326 
Mass. (Dacampia), 326 
Schaer. (Lecidea), 326 
hyalina Berk. (Badhamia) , 306 
Hymenobia Nyl., 316 
Hymenobolina Zukal, 306 
Hymenobolus Zukal, 306 
Hyphelia Fr., 329 
Hypogrea Fr., 329 
Hysterium Tode, 330 

Illosporium Mart., 328, 329 
imbricariae Allesch (Coniothyrium), 327 
Keissl. (Lichenococcum), 327 
imponens Leight. (Lecidea), 330 
indigens Am. (Secoliga), 316 
Rehm (Nectria), 316 
iimata Keissl. (Pharcidia form), 321 
Nyl. (Verrucaria), 321 
Sacc. (Pharcidia), 321 
innatuiaHue (Verrucaria), 321 
Zopf (Pharcidia), 321 
Inoderma S. F. Gray, 331 
inquinans Massai. (Nesolechia), 310 
insidiosa Nyl. (Hymenobia), 316 
Sacc. (Nectria), 316 
insita Stirt. (Lecidea), 309 
Vouaux (? Nesolechia), 309 
inspersa Flk. (Lecidea), 313 
Mudd (Dactylospora), 313 
Rehm (Leciographa), 313 
Tul. (Lecidea), 313 

Transactions British Mycological Society 


insulare Mass. (Endocarpon), 318 
Mig. (Dermatocarpon), 318 
insularis Keissl. (Guignardia), 318 
NyL (Lecidea), 309 
intumescens Flot. (L. badia var.), 309 
Magn. (? Nesolechia), 309 
NyL (Lecidea), 309 

Karsghia Krb., 31 1, 312 
Keissleria von Hobl., 316 
HlHasii Rehm (Mycobilimbia), 310 
kuhnii FkL (Fusisporium), 329 
Sacc. (Fusarium), 329 

Laestadia Ainersw.5 318 
Lahmia Krb., 31 1 
lamyi Rich. (Lecidea), 313 
larbalestieri Quel. (Goniophora), 306 
iaricicola NyL (Xylographa), 315 
Sacc. (Leciographa), 313 
Lasionegtria (Sacc,) Cke., 316 
Leight. (Verracaria)j 319 
iaxa Fr. (Gorticium), 306 
lecanodes Berk. (Dialonectria), 316 
Ges. (Nectria)j 316 
lecanorae KeissL (Ascochytula), 327 
Vouaux (Phoma), 327 
Leciographa Mass., 31 313 
leciographoides KeissL (Mycomelaspilea), 31 1 
Vouaux (Melaspilea), 31 1 
leightonii B. & Br. (Nectria), 316 
Sacc. (Galonectria), 316 
leptogicola Cke. Sl Mass. (Hypocrea), 316 
Sacc. (Broomelia), 316 
von Hohn (Keissleria), 316 
von Hohn (Yatesula), 316 
Leptorhaphis Krb., 325 
Leptosphaeria Ges. & DN., 324, 326 
leptostigma NyL (Lecidea), 308 
Sacc. (Nesolechia), 308 
ieucocephala Fr, (Pyrenotliea), 330 
Pers. (Sphaeria), 330 
leucomelaria Mudd (Sphaeria), 326 
Vouaux (Leptosphaeria), 326 
leucomeiarium B. & V. (Tichothecium), 326 
ievantica Rehm (Gloniopsis) , 330 
Libertella Speg. et Rouin., 327 
lichenicola A. L. Sm. & Ramsb. (Discocera), 


DN. (Bertia), 318 

Cke. & Plow- (Psilosphaeria) ,318 

Harm. (Urceolaria form), 315 

Karst. (Sphaeria), 318 

KeissL (Nesolechia), 309 

Linds. (Torula), 328 

P. & S. (Vouauxiella), 328 

Rehm (Nectria), 316 

lichenicolum Berk. & Br. (Sporotrichum), 330 
Keissi. (PLagadostoma), 318, 327 
KeissL (Sirothecium), 328 
Linds, (Coniothecium), 329 
Mass. ( Melittosporium) ,315 
Vouaux (Dactylium), 329 
Vouaux (Spilomium), 329 
Lighenogonium Pet. & Syd., 327 
lichenodes A. L. Sm. (Diplodina), 327 

Lichenophoma KeissL, 328 
lichenum Schroet. (Celidium), 307 
Somm. (Dothidea), 307 
lignyota Fr. (Fatellaria), 312 
Sacc. (Karschia), 312 
Listerella Jahn., 306 
lithina Leight. (Pyrenothea), 330 
Tayl. (Verrucaria), 330 
iongius NyL (Odontotrema), 315 
lopadii Anzi (Celidium), 324 
Zopf (Merismatium), 324 
Lophothelium Stirt., 319 
lutea Leight. (Pyrenothea), 330 

Magrosporium Fr., 329 
macvicari Grelet (Stenocybe var.), 314 
major NyL (Stenocybe), 314 
majus Leight, (Odontotrema), 315 
margacea Leight., 1851 (Verrucaria), 323 
Massaria DN., 326 
Massariella Speg., 3.26, . 
meialeucum Ach. (Spiloma), 328 
Me'lanomma Nke. & FkL, 325 
melanospermum Macbride (Didymium), 306 
meianospora Hepp. (Didymosphaeria), 326 
melaspermeila NyL (Arthonia), 312 
KIelaspilea NyL , 3 1 1 , 314, 315 
melaspora TayL (Verrucaria), 319 
Melittiosporium Corda, 315 
Merismatium Zopf, 324 
Metasphaeria Sacc., 316 
microcephala Nyi, (Sphinctrina), 314 
Microce'ra Desm., 328 
micropliorus NyL (Endococcus), 321 
microspermus Linds. (Abrothaiius var.), 312 
TuL (Abrothaiius), . 312,, 
microspila Krb. (Artliopyrenia), 322 
Wint. (Pharcidia), 322 . , 

microsticta' Mudd (Acarospora var.), 326 
microstictica Leight. (Verrucaria), 326 
Wint. (Didymosphaeria), 326 ' 

Vain. (Didymosphaeria), 326 
micros tic ticum Leight. (Endocarpon), 326' 
Microthelia Krb., 318, 320, 321, 323, 324, 
326, 327 

micula Flot. (Microthelia), 326 
miliaris Walk. (Peziza), 309, 
minus NyL (Odontotrema), 315 
minutum Fr. (Graterium), 306 
mollis Leight. (Pyrenothea), 330 . 

Tayl, (Verrucaria), 330 
Mollisia Fr,, 308 
M0LLISIEL.LA Boud., 308 
moorei Linds. (Abrothaiius), 312, 328 . 
moriformis Th. Fr, (Biatorella), 314 
Mullerella, H epp, 317 
Mycobagidia' R ehm, 311,314',' 
Myggbilimbia R'Chm, 310'' ^ 
Mycomelaspilla Reinke, ,3 1 1 , 
Mycosphaerella JO'hans.,.'3i9 ' 

Myriangium M, & B., 318 
Myxogocgus 'Thaxt., 306 ■ ,,, 

Nectria Fr., 316, 317 
Nectriella Nitsch, 316 


British Fungi Parasitic on Lichens. W. Watson 

neglecta Hue (Grocynia), 309 
Nyl. (Lecidea), 309 
Vain. (Nesolechia), 309 
neglectum Desm. (Epicoccum), 329 
Neolamya Theiss. & Syd., 327 
neottizans A. L. Sm. (Didymosphaeria), 326 
Leight. (Verrucaria), 326 
Zopf (Leptosphaeria) , 326 
nephromaria Nyl. (Arthonia), 308 
Nesolechia Mass., 308, 314 
nigritella A. L. Sm. (Poiyblastia), 324 
Am, (Poiyblastia), 324 
Nyl. (Vermcaria), 324 
Vouaux (Merismatium), 324 
nigropunctata Hook. (Lecidea), 326 
liigmm Lam. (Goniothecium), 328 
niveoatra Borr. (Verrucaria), 330 
Leight. (Pyrenothea), 33a 
notarisii ZahL (Gyplielium), 329 

Obryzum Wailr., 319 
obscurata A. L. Sm. (Bilimbia var.), 31 1 
Rehm (Mycobilimbia), 310 
Somm. (Lecidea van), 310 
ZahL (Bacidia), 310 
obscuroides Linds. (Lecidea), 309 
octospora Wats. (Microthelia form), 323 
Odontotrema Nyl., 315 
olivaceum DC. (Goniothecium), 328 
Opegrapha Humb., 330 
opegraphicola Linds. (Microthelia), 317 
Zopf ( Mullerella) ,317 
Orbicula Cke., 316 
Orbilia Fr., 308 

oropensis Hohn. (Ciliomyces), 317 
oxyspora Mass. (Nesolechia), 309 
Nyl. (Lecidea), 309 
oxyspoms TuL (Abrothallus), 309 

pallens Nyl. (Xylographa var.), 315 
papiliosa Ach. (Pertusaria var.), 329 
paradoxa Jahn (Listereila), 306 
Paralaestadia Vain., 318 
parallela Fors. (Xylographa), 315 
Faranectria Sacc., 317 
parasemoides Am, (Celidium form), 307 
Nyl. (Arthonia), 307 

parasitica Arn. (Phaeospora), 322, 323, 324 
Mass. (Leciographa), 313 
parasiticus Sm, (Lichen), 312 
ZahL (Diploschistes var.), 315 
Zuk. (Hymenobolus), 306 
parellaria A. L. Sm. (Leciographa var.), 313 
Nyl. (Lecidea), 313 
Sacc. & D. Sacc. (Leciographa), 313 
parietinaria Linds. (Mkrodielia), 320 
parmeliarum Gke. (Heptameria), 324 
' Gke., (Melanomma), 324 
Cke. & Plow. (Psiiosphaeria), 324 
Oliv., (Bueliia),''3i2': 

'Phil. & Plowr.' (Sphaeria), 324 
Sacc. (Leptosphaeria),. 324 
Somm (Lecidea) ,3 1 2 
Vouaux (Abrothallus), 312 
Vouaux (Phaeospora), 324 

parmularia Berk. (Sphaeria), 319 
Leight. (Astrothelium), 319 
Sacc. (Vaisaria), 319 
particular^ A. L- Sm. (Bueilia), 312 
Nyl, (Lecidea), 312 
patersoni Stirt. (Melaspilea), 315 
peltigerae KeissL (Libertelia), 327 
Lib. (Zythia), 327 
Mont. (Sphaeria), 327 
Phil. & Plowr. (Nectria), 316 
Th. & Syd. (Neolamya), 327 
peltigerea Rehm (Scutiila), 308, 310 
Th. Fr. (Arthonia), 308, 310 
pelvetii Arn. (Conida), 308 
Cke. (Homostegia), 308, 317 
peripherica Arn. (Phaeospora), 323 
Mudd. (Microthelia), 323 
Tayl. (Verrucaria), 323 
periphericus Gromb. (Endococcus), 323 
perpusiiia Leight. (Verrucaria), 320 
perpusillum Am. (Tichothecium), 320 
perpusillus Nyl. (Endococcus), 320 
persimilis Nyl. (Lecidea), 313 
petraeicola Linds. (Microthelia), 323 
Peziza Dill., 308 
Pezizella Starb., 308 
Phacopsis TuL, 307, 308 
Phaeospora Hepp., 323 
Pharcidia Krb., 321, 322 
Phoma Fr., 327 

Phragmothyrium Hdhn, 316, 325 
Phymatopsis Linds. ,312 
physciae Kalch. (Gymnosporium), 328 
Pers. Fr. (Aegerita), 328 
Sacc. (Coniosporium), 328 
physciicolum KeissL (Discothecium var.), 320 
Nyl. (Mycoporum), 320 
Vouaux (Discothecium), 320 
piggotii Berk. & Br, (Dothidea), 317 
Karst. (Homostegia), 317 
pinicola Th. Fr, (Biatorella), 314 
pituphloia Leight. (Verrucaria), 325 
Pleospora Rabh., 326 
plerospora A. L. Sm. (Lecidea), 309 
pleurocarpa A. L. Sm. (Lobaria var.), 307 
plumbina Anzi (Leciographa), 31 1 
Nyl. (Lecidea), 31 1 
KeissL (Lahmia), 311 
Vouaux (Mycobacidia), 311 
podetiicola KeissL (Dendrophoma), 327 
Polyblastia Mass., 326 
PoLYCEPHALUM Kalchb. & Gke., 306 
polyspora Hepp (Mullerella), 317 
Psilosphaeria Cke., 316, 318, 324 
Psora Haller, 316 

psoromoides Borr. (Vermcaria), 318 
Hook. (Endocarpon), 318 
KeissL (Guignardia), 318 
Vouaux (? Laestadia), 318 
Wint. (Physalospora), 318 
puHcare Pers. (Hysterium), 330 
pulmonacea Ach. (Sticta), 307 
pulmonaria Hoffm. (Lobaria), 307 
pulposa Leight (Verrucaria), 330 
pulposi Vouaux (Didymella), 325 
Zopf (Didymosphaeria), 325 



338 Transactions British Mycologtcal Soaety 

puncta MassaL (Nesoiechia), 31 1 
punctilla Nyl. (Arthonia), 308 
Vouaux (Gonida), 308 
Wint. (Pharcidia), 322 
pycnostigma Nyl. (Vermcaria), 326 
Sacc. & D. Sacc. (Leptosphaeria), 326 
pygmaea Krb. (Microtheiia), 320 
pygmaeum Krb. (Tichothecium), 320 
Pyrenogoggus Wheld. & Wils., 323 
pyriformis A. L. Sm. (Myxococcus), 306 

ramalinae Oliv. (Spiloinium), 328 
Raphiospora Mass. (Bacidia with carbonaceous 
apothecia), 31 1 

resinae Mudd. (Biatorella), 314 
Th. Fr. (Biatorella), 314 
Rhagadostoma Krb., 318 
Rhizoctonia DC. ex Fry, 307 
rhyponta Borr. (Verrucaria), 322 
Mudd (Arthopyrenia), 322 
rhytidiosporum Berk. & Br. (Stilbum), 306 
rimosicola Leight. (Verrucaria), 323 
Mudd (Microtheiia), 323 
Zopf. (Phaeospora), 323 
rimosicolum Arn. (TicliotJiecium), 323 
robergei Mont. & Desm. (Nectria), 316 
Weese (Nectriella), 316 
rosea Fr. (Hyphelia), 329 
roseum Berk. (Dactylium), 330 
Fr. (Hyphoderma), 329 
Link. (Trichothecium), 330 
Mart. (Illosporium), 329 
rudis Borr. (Verrucaria), 330 
Leight. (Pyrenothea), 330 
rufum Fr. (Agyrium), 307 
rugulosa Borr. (Verrucaria), 319 
Mudd (Microtheiia), 319 
rugulosus Nyl. (Endococcus), 319 
ruiziana MiilL-Arg. (Graphina), 330 

saxatilis Krb. (Bueilia), 312 
Rabh. (Trachylia), 312 
Rehm (Karschia), 312 
Schaer. (Calicium), 312 
scabrosa Krb. (Bueilia), 312 
Rehm. (Karschia), 312 
scapanaria Carrington (Lecidea), 313 
A. L. Sm. (Leciographa), 312 
scaphoidea Stirt, (Xylographa), 315 
ScHizoxYLON Pers., 315 
scoriadea Cke. (Massaria), 326 
Fr. (Sphaeria), 326 
Sacc. (Massarieiia), 326 
ScuTULA TuL, 310 
Scutularia Karst., 314 
Secoliga Stitzb. ,316 
silacea Fee (Gassicurtia), 329 
silaceum KeissL (Coniothecium), 329- 
Nyl. (Spilomium) , 329 
SiROTHEGiUM Karst., 328 
smithii Linds. (Lecidea), 312 
Tul. (Abrothellus), 312 
solorinaria Linds. (Microtheiia), 318, 327 
Vouaux (Diplodina), 318, 327 
Sorotheua Kib., 320 
Speggazinia Sacc., 329 

sphaerale Fr, (Sclerococcum), 329 
Keissl. (Goniothecium), 329 
Sphaerella Ges. & DN., 321 
Sphaeria Hall, 321, 322, 323, 324, 331 
sphaerica Schaer. (Lecidea), 326 
Sphaerostilbe Tul., 328 
Sphaerulina Sacc., 318, 319 
Sphingtrina Fr., 314 

sphinctrinoides Beri. & Vogi. (Didymeila), 325 
Spiloma auc. ant., 328 
Spilomela Keissl., 329 
Spilomium NyL, 329 
spiiomatica Th. Fr. (Xylographa), 315 
Sporotrighum Link., 330 
squamaricolum Linds. (Gelidium), 307 
squamarioides Keissl. (Discothecium), 320 
Mudd (Sphaeria), 320 
Wint. (Tichothecium), 320 
Zopf (Sorothelia), 320 
Stegia Fr., 324 
Stenocybe NyL, 314 
stereocauHcola Linds. (Microtheiia), 320 
stereocaulicolum Vouaux (Discothecium), 320 
stereocaulina Keissl. (Gonida form), 308 
Ohl. (Arthonia var.), 308 
stereocaulorum Krb. (Scutula), 320 
Nyl. (Lecidea), 320 
Th. Fr. (Biatorina), 320 
Sticta Schreb., 307 
stictarum Tul. (Gelidium), 307 
DN. (Sphaeria), 307 
stictica Fr. (Pyrenothea), 330 
stigma Krb. (Tichothecium), 320 
Zopf (Discothecium), 320 
Stigmatella L6v., 306 
stigonellum Schaer. (Calicium), 313 
Zahl. (Gypheiium), 329 
Stilbum Tode, 306 
Strigula Fr,, 316 

subfuscicola Linds. (Microtheiia), 308 
subvarians Nyl. (Arthonia), 308 
sulphurea Leight. (Pyrenothea), 330 
supernuia A. L. Sm. (Biatorina), 310 
Nyl. (Lecidea), 310 
superposita Nyl. (Verrucaria), 322 
Sacc. & D. Sacc. (Pharcidia), 322 
superpositum A. L. Sm,. (Theiidmm), 322 
supersparsa Arn. (Phaeospora), 324 
Keissl. (NesolecMa var.), 3.10 ■ 

Nyl. (Lecidea), 310 

talcophila Kxb, (Karschi.a), 31,2 
tartaricola Cke. (Orbicula), 316 
Linds. (Verrucaria), 326 
Nyl. (Sphaeria), 316 
tartarina Keissl (Metasphaeria), 325 
Nyl. (Verrucaria), 325, 
tenuis Nees ex Wallr., (Altemaria), ,329 
thalamitus Nyl , (Endococcus) 32 1 
thallina Cke. (Sphaerella), 321 
Sacc. (Epicymatia), 321 ■ 
thalloides Rehm (Karschia), 312 ' 
thallophila Cke. (,Sphae.rella),^322 (Sphaeria),' 322 
'Sacc. (Epicymatia), 322 
Stevenson (Psilosphaeria), 322'' 


British Fungi Parasitic on Lichens. W. Watson 

Thelidium Mass., 321, 322 

Thelocarpon Nyi., 317 

Thrombium Wal!r., 331 

Tighothegium Fiot., 320, 321, 326 

Torula Pers. Fr., 328 

Traghylia Fr., 312 

Trichothegium Link, 330 

triphractoides A. L. Sm. (PPharcidia), 322, 


Leight. (Verrucaria), 322 
Nyi. (Endococcus) 5 322, 323 
tubaeformis Massal. (Sphinctrina), 314 
turbinata Fr. (Sphinctrina), 314 

uncialicola Vouaux (Phoma), 312, 328 
Zopf (Phyilosticta), 328 
usneae Jatta (Buellia), 312 
Rabh, (Abrothalius), 312 
Rehm (Abrothalius form), 312 
utricularis Berk. (Badhamia), 306 

Valsaria Ces. & DN., 319 
varians Arn. (Celidium), 307 
Hall (Lichen), 307 
Nyi. (Arthonia), 307 
variolosum T. & B. (Spiloma), 328 
varium Massal. (Celidium), 307 
Tul. (Phacopsis), 307 
ventosaria Linds. (Sphaeria), 321 
ventosicola Leight. (Verrucaria), 321 
Mudd (Microtheiia), 321 
ventosicolum Wint. (Tichothecium var.), 

ventosus NyL (Endococcus), 321 
vermiceliifera Leight. (Pyrenothea), 330 
vermicuiaria Linds. (Microtheiia), 320 

vermicularis Arn. (Nesolechia), 314 
Gray (Gerania), 320 
Keissl. (Stegia), 314 

vermicularium Vouaux (Discothecium), 320 
vermifera Keissl. (Spilomela), 314 
Leight. (Melaspilea), 314 
Sacc. (Scutularia), 314 
Vouaux (Mycobacidia), 314 
vermiformis Leight. (Melaspilea), 314 
Verrugaria Pers., 317-^26 
verrucariae (Didymella var.) (Zopf) Vouaux, 


verrucosaria Linds. (Arthopyrenia), 325 
versicolor Lister (Badhamia), 306 
Smith (Spiloma), 328 
vesicularia Arn. (Phaeospora), 324 
Linds. (Microtheiia), 324 
viridescens (Fr.) Biatora, 306 
viteilinaria NyL (Lecidea), 310 
Rehm. (Nesolechia), 310 
vouauxi B. de Lesd. (Diplodina), 327 
Vouauxiella Pet. & Syd., 328 
vugasus Eng. Bot. (Lichen), 330 
vulgaris Fkl. (Epicymatia), 322 
Mass. (Sphaerella), 322 

walirothii Nyi. (Lecidea), 310 
watsoni Rilstone (Gloniopsis), 330 
welwitzschia Mont. (Abrothalius), 312 
Tul. (Abrothalius), 312 

Xylographa Fr., 314, 315 

Yatesula Syd., 316 

zwackhii Cromb. non Massal. (Lecidea), 313 
Zythia Syd., 327 

{Accepted for publication 17 March 1947 ) 


[ 340 ] 


'By a Sub-committee* o.f the Plant Pathology Committee 
OF THE British Mygologigal Society 

The third edition of the List of Common British Plant Diseases, compiled by 
the Plant Pathology Committee of the British Mycological Society, was 
published in 1944. Since then a number of emendations and ad^tions 
have become necessary, and these are given below, together with the 
number of the page in the third edition where the change or addition 
should be made. Following the practice now general among botanists, 
specific epithets have been decapitalized. Until there is a generally accepted 
set of international names for viruses, the common names used for virus 
nomenclature in the Review of Applied Mycologp (Special Part, vol. xxrv, 
part 13, 1946) are accepted. 

Apple, p. 9. To Bitter Rot add ‘and Gloeosporiumperennans Zeller & Childs’. 
Apple, pp. 9-1 1. Add the four entries: 

‘Dry Eye Rot and Soft Rot. . .Botrytis cinerea Fr. 

‘Perennial Canker. . .Gloeosporium perennans Zeller & Childs, 
‘Chlorosis (lime-induced) . . .Iron deficiency, 

‘Magnesium Deficiency. . .Non-parasitic.’ 

Barley, p. 12. Brown Rust. The nzmc Puccinia hordei Otth (1871) was 
considered in 1939 (these Transactions, xxiii, 279) and, though it was 
recognized to be the earliest name, it was at that time thought best to 
regard it as a nomen ambiguum. However, Buchwald (1943) has restated 
his case, and has now provided a valid name for the homonym 
P. hordei Fuckel (1873). A hordei Otth, which is the valid name under 
the Rules for this Brown Rust fungus, has recently been accepted by 
the U.S. Department of Agriculture (Stevenson & Johnson, 1946) and 
we recommend that it be accepted here also. In view of the confusion 
in the literature over the names P. anomala and P. simplex, it is hoped 
that P. hordei Otth may provide a name to be used everywhere. 
Bean, Broad and Field, p. 13. Chocolate Spot. To Botrytis cimrea Fr. add 
dLikd. B. fabae Sardina’. 

Cherry, p.^; 17. To Bacterial .Cankef add ‘and Leaf Spot’, The same 
addition under Plum, p. 32. 

Gugumber, p. 19. Add the entry ‘Stem and Fruit Rot. .^Fythmmultimum 
Trow SiXid P. apkanidermatum (Eidson) FkzpdLtnck\ 

* The .sub-committee is G. R. Bisby (convenor), K. St G. Cartwright, Cro,xa!I, 
E. W. Mason, W, G. Moore and Miss E. :M. Wakefield. These emendations have been 
approved by the Plant Pathology Committee. 

Third Edition of the List of Common British Plant Diseases 341 

Flax, p. 21. To Fusarium Uni should be added the synonym F. oxysporum Fr. 
f. Uni (Bolley) Snyder & Hansen, Although the proposals of Snyder 
and Hansen regarding formae of F. oxysporum have been commonly 
accepted in North America, it seems best to follow Wollenweber and 
Reinking until there is more general agreement as to the taxonomy 
and nomenclature of Fusarium. After Rust add the entry ‘Mildew. . . 
Oidium Uni Skoric’. 

Gladiolus, p. 22. Botrytis Rot and Core Rot. Change ‘Botrytis sp.’ to 
^ Botrytis cimreaFtd 

Grasses, p. 23. The entry ‘Stripe Smut. . .Ustilago striiformis (Westend.) 
NiessF requires no change; but, to lessen the danger that someone 
might propose an undesirable change, we endorse the view presented 
by Stevenson (1946) that the earlier name U. salvei B. & Br. should 
be considered a nomen ambiguum. 

Mushroom, p. 27. Add the entry ‘Verticillium Disease. . .VerticilUum 
malthousei Ware’. 

Oak, p. 27. Mildew. Robertson and Macfarlane (1947) report a few 
perithecia, which they identified as Microsphaera alphitoides Griff. & 
Maubl., on Quercus robur in Herts. 

Oats, p. 28. Loose Smut. Stevenson & Johnson (1944) find that the 
fungus should be cited as Ustilago avenae (Pers.) Rostr. (1890). Jensen 
in 1889 used the binomial U. avenae as a nomen nudum. 

Onion, p. 28. Add the entry ‘Leaf Blotch. . . Heterosporium allii Ell. & 
Martin var. cepivorum Nicolas & Aggery’. 

Plum, p. 32. Add the entry ‘Chlorosis (lime-induced). . .Iron deficiency’. 
Potato, p. 34. Add the three entries : 

‘Leaf Scorch. . .Potassium deficiency and other causes. 

‘Premature Tuber Formation (Little Potato) . . . Non-parasitic. 
‘Glassiness. . .Non-parasitic.’ 

Rhododendron, p. 35. To Leaf Scorch add ‘and Bud Blight’. 

Spinach, p. 37. Add the entry ‘Leaf Spot. . .Heterosporium variabite Cooke’. 
Strawberry, p. 37. Add the entry ‘Leaf Blotch.. . .Zjthia fragariae 
Laibach. The perfect state is an undetermined species of Gnomoniah 
Tomato, p. 39. To Grey Mould and Botiytis Fruit Spot add ‘and Soft Rot’. 
Tomato, pp. 39-40. Add the three entries : 

‘Alternaria Blight. . .Altemaria solani (Ell. & Martin) Sor. 

‘Bacterial Canker. . . Corynebacterium michiganense (E. F. Sm.) H. L.Jens, 
‘Magnesium Deficiency. . .Non-parasitic.’ 

Turf, p. 40. Fetch (1945) has strongly reinforced the view expressed on 
p. 7 of the List that ‘ It might have been even better to accept Calonec- 
tria nivalis Schaffnit’ instead of C. graminicola Wollenw., and it is 

therefore proposed that C. KjWir be accepted. 

Viola, p. 41. Change the first entry to read: ‘Root and Stem Rot.. . 
Pytkium violae Chesters & B.ic)svaa.n, Myrothecium roridum Fr., and 
sometimes other fungi.’ 

Walnut, p. 42. Graft Disease. Delete the entry regarding the perfect 
State. Baker and Thomas (1946) did not obtain any evidence that 
Chalaropsis thielavioides has a. ^exiect sta.te. 

342 Transactions British Mycological Society 

Wheat, p, 42. Add the entry ‘ Sharp Eyespot . . . Corticium solani (Prill. & 
Delacr.) Bourd. & Galz., stat. mycel. Rhizoctonia solani Kiihn’. 

The following names are now to be added to the Hst of author’s names : 
Aggery [Berthe] Nicolas [G.] 

Chesters [C. G. G.] Sardina [J. R.] 

Childs [L.] Skoric [V.] 

Jens[en] H. L. . Ware [W. M.] 

Laibach [F.] Zeller [S. M.] 

Martin [G., 1827-86] 

The following dates can be added to the printed list: 

Henn[ing, E., 1857-1929] 

Jones, L. R. [1864-1945] 

Rabat [J. E., 1849-1925] 

King [Charlotte M., 1864-1937] 
Kj[eb[ahn, H., 1859-1942] 
Lambert, F. [1859-1940] 

Lind [J., 1874-1939] 
Schilb[ersky, K., 1863-1935] 
Syd[ow, H., 1879-1946] 
Tak[ahashi, Y., ?-i9i4] 

Towns [end, C. O., 1863-1937] 


Baker, K. F. & Thomas, H. E. (1946). Failure of bud and graft unions of rose induced 
by Chalaropsis thielavioides. Phytopathology, xxxvi, 281-91. 

Buchwald, N. F. (1943). liber Pucdnia hordei Otth (Syn. P. simplex (Kcke.) Erikss. & 
Henn.) und P. hordei-murini n.n. (Syn. P. hordei FckL). Ann. mycoL, Berl. xli, 306-16. 
Fetch, T. (1945). Additional notes on British Hypocreales. Trans. Brit. myc. Soc. xxvn, 


Robertson, N. & Macfarlane, L (1947). The occurrence of perithecia of the oak mildew 
in Britain. Trans. Brit. myc. Soc. xxix, 219-20. 

Stevenson, J. A. (1946). A nomenclatorial discussion of Ustilago striiformis. Plant Dis. 
Reptr. XXX, 57. 

Stevenson, J. A. & Johnson, A. G. (1944). The nomenclature of the cereal smut fungi. 
Plant Dis. Reptr. xxvni, 663--70. 

Stevenson, J. A., & Johnson, A. G. (1946). The nomenclature of the barley leaf rust. 
Plant Dis. Reptr. xxx, 372. 

{Accepted for publication i8 August 1947) 

[ 343 ] 


In 1941 the Society’s Plant Pathology Committee formed a Sub-Committee 
to study methods of disease measurement in the field. A preliminary report 
was published (Moore, 1943) and this was followed by short progress 
reports {Trans. 1943-7). Altogether, methods of measuring disease in- 
tensity have been laid down for the following diseases : 



Sugar Beet 

Smuts; Take-all; Eyespot; Brown Foot Rot. 

Blight; Leaf Roll; Leaf Drop Streak; Rugose Mosaic 
( = Severe Mosaic). 



The methods recommended for all these diseases, except Apple Scab, 
have been given in the progress reports. 

Percentage infection method 

The simplest method of disease measurement is to estimate the percentage 
infection. For wide surveys the records may take the form of the percentage 
of crops in a district in which a disease occurs, and may be designated as P 
(disease prevalence) . 

Usually the figure obtained for the amount of disease is the percentage 
of plants infected in a whole plot, or in samples taken from the crop at 
random, and is obtained simply by counting and averaging the samples. 
This method is applicable to all the diseases in the above list, except Potato 
Blight and Apple Scab, which normally develop on every plant in the crop. 
Figures obtained in this way may be designated by the letter I (percentage 
of infection) . 

The method of taking the samples will vary with the crop and the 
disease. To save time and labour it is desirable to take no more than the 
minimum required to give accurate results. Generally speaking ten samples 
are adequate within one plot or crop, provided they are satisfactorily 
randomized. If, however, there is considerable variation in the results 
from the ten samples, another ten samples should be taken. With plots in 
which varieties are replicated several times, the random sampling should 
be made over the whole of the replicates, and not concentrated on a few 
plots with only a proportion of the replicates sampled. 

Disease index method 

For diseases in which the amount of disease is variable on different plants 
within the crop the figure for I may give an inadequate measure of the 
disease intensity, and it becomes desirable to assess the arnount of disease 
on each plant. This has been done by reference to an arbitrary scale. 

The simplest type of scale is of the qualitative type, e.g. healthy, slight, 
severe. But even this scale has drawbacks: (i) different observers, or the 

344 Transactions British Mycological Society 

same observer in different seasons, may show disagreement over borderline 
cases, and (2) comparison of disease severity may be ambiguous, e.g. it is 
not certain whether 50 % slight and io % severe is more or less severe than 
30 % slight and 15 % severe. The first difficulty, or the personal equation, 
may be largely surmounted where groups of observers work together to 
check each other’s assessments periodically, but the second becomes 
important as more elaborate scales are adopted. 

Americans have widely used the ‘Disease Index’ method by which the 
observations of disease intensity are averaged out to a single figure (Walker, 
Larson & Albert, 1938). Suppose there are five grades i, 2, 3, 4, 5 
measuring the amount of disease : if JV is the total number of plants, and 
jVi, jV2, jV3, jV4, Ws are the numbers of plants in each grade, then the 
Disease Index figure, which may be designated DI, is 

JVi+2jV2 + 3W3+4W4 + 5 jV 5 ^^ 

.... _ . .... ^ J 00# 

Any inaccuracies in placing the observations in the right categories will 
increase the error of the final figure, and another error will be introduced 
if the amounts of disease in the different categories are not proportional to 
the numbers of the categories. A few observers have surmounted the 
second difficulty by giving numbers to the categories equal or proportional 
to the percentage of leaf area destroyed. This modification leads on to a 
third method of disease measurement. 

Percentage affected leaf-area method 

Direct estimation of the percentage leaf area affected giv^ figures 
which may be designated DM. 

The method is particularly appropriate for diseases such as Potato Blight 
in which normally every plant is affected. Some sort of standard' is 
essential, although the amount of use of, or frequency of reference to, the 
standard will vary with individual skill and temperament. For field survey 
it is usually sufficient to classify into ‘group percentages' with a range of 
5 to IO, or even up to 25 at the less important parts of the scale. Some 
prefer to use ‘standard area diagrams' or ‘standard photographs', but the 
method which is gaining most support in this country at present is the use 
of the ‘standard (descriptive) key', carefully prepared after preliminary 
investigations of the disease, and modified during the first few years of its 
use in the light of field experience. 

With the aid of such standard keys the percentage leaf area affected by 
the disease can be estimated fairly accurately. The assessments can be 
made on single plants, but are just as easily carried out on small handfuls 
selected at random, or even on whole plots, for which the method is 
primarily designed. With certain crops it may be desirable to record the 
observations for two or three different height levels (^, m and bottom, 
middle and top). 

Experience has shown that the personal equation is less serious with tliis 
than with other methods, because the measurements have reference to 

Disease Measurement in Plant Pathology 345 

a standard key. Also the values for DM have a direct biological meanings 
and observations on different dates on the same plots or fields in any 
season can be plotted in a curve, which is normally sigmoid in character. 
Furthermore, it is an advantage to have one simple uniform method for 
different diseases. It is, of course, necessary to construct a separate key 
for each disease, and this is a matter for specific investigation. It is a 
further advantage that, as the crop yield depends on the amount of leaf 
functioning during the season, it should be possible to correlate disease 
measurement with crop yield. Estimates of losses of crops, and decreases 
in financial returns, with resulting wise direction of national research into 
control methods for the diseases placed in order of priority, should be the 
ultimate objective of all disease measurement work. 

A good example of the value of these methods is given by the observations 
(Large, 1945) on potato blight spraying trials. 

Enough of the preliminary work has been done to enable rapid progress 
to be made, even if that progress is along a rather irregular front. A lead 
has been given and interest in the subject has been created in this country. 
The Society’s Sub-Committee is therefore confidently content to leave the 
future organization of disease measurement work in the hands of the 
Disease Assessment Sub-Committee recently appointed by the Conference 
of Plant Pathologists of the National Agricultural Advisory Service. It is 
intended that this new Sub-Committee shall be broadened to include 
workers interested primarily in the research side as well as the specialist 


Large, E. G. (1945)* Field trials of copper fungicides for the control of potato blight. I. 

Foliage protection and yield. Ann, appL Biol, xxxn, 319-29. 

Moore, W. C. (1943)* The measurement of plant diseases in the field. Trans, Brit, myc, 
Soc, XXVI, 28-35. 

Tram, (1943-7). The measurement of plant diseases in the field. Tram, Brit, myc, Soc, 
XXVI, 1943, 172-3 and XXXI, 1947, 140-41. 

Walker, J. C., Larson, R. H. & Albert, A. R. (1938). Studies of resistance to potato 
Scab in Wisconsin. Amer, Potato J, xv, 246-52. 

{Accepted for publication 30 June 1947) 

1 346 ] 



The publication of mycological exsiccatae in the past has been almost exclusively a matter 
of private initiative. The great value of this tremendous collecting work performed during 
the past century should not be underrated, but it must be admitted that in most cases the 
material has been collected by chance and determined by mycologists who could not 
possibly be specialists in the numerous fungous groups with which they dealt. The many 
misidentified specimens do more to complicate than to simplify identification of new 
material, and a short cut to sure determination is badly needed, especially for plant 

The publishing of an International Herbarium, based on contributions from selected 
specialists, with the nomenclature steadily kept up to date, would make it possible to 
establish a much needed co-operation between the mycological taxonomists and the plant 
pathologists, who often cannot spend much time on taxonomic studies. I propose the 
following principles to be taken into consideration for the realization of the suggested 

I. Collecting 

The purpose of the herbarium is to facilitate identification of plant pathogenic fungi: 
hence, saprophytic forms belonging to genera containing parasitic forms must be included. 
Examples of groups entitled to be included are: Alternaria^ Ascochyta, Boiryiis, Cercosporay 
Ciboria, Cladosporiurriy Colletotrichuniy Coniothyrium^ Cyitndrocarpon, Cystopusy Cucurbitariay 
Didymellay Diplocarpony Entylomay Erysiphaceae, Fusariuruy Gloeosporiumy Helmintkosporiumy 
Heterosporiumy Marssoninay Mycogone, Mycospbaerellay Nectriay Oimlaria, Peromsporay Pesta-^ 
lozzicCy Phorm, PhyllostictUy Phyiophthoray Plasmodiopkoray PolyporuSy PythiuMy Ramuiaridy 
Rhizoctonidy Rhytisma, Sclerotinidy Septoria, Sphaceiorm, Stagonosporuy Stemphylmny Taphrina* 
ceae, Uredinales, Ustilaginales, Venturidy Vermiculariay VerticUlium. 

Whenever possible material of the same fungus (same collection) should be issued 
from (i) a natural substratum, (2) an artificial medium, as an isolate. 

All the material for a given specimen should be taken from one locality and one crop 
or substratum. Enough material should be collected to satisfy any request to be expected 
for a long period of time. If, for example, 500 labelled packets should be made to cover 
the demand for some years, an ample supply of I'eserve material should be kept for 
dividing and packeting later, if that should prove necessary. 

2. Determination 

The determination should be based on the co-operation of specialists, invited to take care 
of the groups in which they have specialized. More than one specialist in a group can be 
invited, and it is even preferable that the material be determined by two specialists, who 
should be cited as determiners on the label. For a long time it would probably be 
‘itf impossible to find specialists for all the fungus groups that should be represented. But 

in any case it would be preferable to limit the issuance of items to the forms for which 
^ a well-documented determination could be given. Specialists of single species could be 

invited to contribute material of the species in question. 

^ In brief, the dd herbaria rendered material incidentally found and more or less tenta- 
tively determined. For the suggested herbariuniy the collecting work should follow a pre- 
arranged plan: the forms to be included should be well known to specialists and should 
— >as far as possible — ^be chosen in advance and, at the beginning, be selected preferably 
from the economically or taxonomically more important forms. 

Though pleased to publish this contribution from an eminent Danish colleague, the 
Council of the Society wish to point out that they do not necessarily share his views. 

International Herbarium of Plant Pathogenic Fungi 347 

3. Labelling 

The following data are proposed to be given on the label (see sample label below) : 

{a) Name of the exsiccata 

{b) Issue number of the specimen 

[c) Name of the fungus, including author(s) and date of publication 

(d) Synonyms, including author’s name (s) followed by date of publication 
{e) Substratum (host, artificial medium) 

{/) Date and locality of collection. For material obtained by culture : 

(aa) Date of isolation 

{bb) Substratum from which the fungus is isolated and the date and locality of 
the collection of that material 

{cc) Date of transferring the fungus to the medium used for the issue 
{dd) Date of drying the culture 
{g) Bibliography: 

{aa) Reference to original diagnosis 

{bb) Reference to the first use of accepted name 

{cc) Reference to diagnosis in Saccardo Sylloge Fmmmm 

{dd) Monograph (s) 

{ee) Most important literature. If data on the specimen are published, this 
should be specially indicated 
(A) Name of determiner (s) 

{i) Remarks on possible deviations of the specimen from the normal form (the type) 
(j) In case of dispute on the nomenclature, references to be made to papers containing 
the determiner’s arguments. Where there is complete disagreement between two 
selected authorities (as to type of the species, etc.) duplicate labels should be 

{k) Date of issuance of the label, and number of the edition, e.g. March 1952, 2nd edition 

No. 123 1st edition of label, issued Aug. 1952 

Heterosporium iridis (Faut. & Roum.) Jacques 1941 
Syn. : Scolecotrichum iridis Faut. & Roum. 1891. 

Heterosporium montenegrinum Bub. 1903. 

Scolecotrichum cladosporoideum Maire 1906. 

On malt extract agar, isolated 19 Feb, Iris germanica (coll. 2 September, 

1949 in Branchport, New York, U.S.A. — material issued as Herb. Phytopath. N. N. 

No. 122)3 transferred last time 22 March, 1950, the culture dried 3 April, 1950. 

References : 

Original diagnosis', Roumegudre: Fung. Sei. Gal. Exs. No. 5689, 1891. 

New combination : Jacques, J. fimile : Studies in the genus Heterosporium, — Contr, de 
rinsL Bot, de r Unit}, Montreal, No. 39: 1--59. 1941* 

Saccardo: Sylloge Fungorum 10: 600. 1892. 

Klebahn, H. : tiber drei auf Iris gefundene Perithezien und die zugehorigen 
Konidienpilze, — Ber, Deut, Bot, Gesell, 42 (General versammlungs-Heft) : 60-7 1 . 

Perrault, G. : A common leaf spot of Iris in Quebec. — Quebec Soc, Prot. Plants 
Ann, Rep. ig: Sy-iog, 1927. 

Ramsbottom, J. K. : Iris leaf-blotch disease {Heterosporium gracile Sacc.) — J , Roy, 
iSbi;. (London) 40: 481-92. 1915. 

SiBiLiA, G. ; ‘Saltazioni’ in Heterosporium gracile, — Boll, Staz- Put. Veg, Romuy 
N.S. ' 14:. 447-474. 1934. 

Tisdale, W, B.: Iris leaf spot caused by Didymellina iridis,— Phytopath, 10: 
148-163. 1920. 

Det. N. N., Montreal, Canada, and X. X., London, England. 

Transactions British Mycological Society 

New labels should be issued to provide for corrections of possible errors in identifications, 

toVate care of new combinations, new synonyms, or to cite new important literature-m 
brief, to keep the herbarium up-to-date. 

4 Publication of new systematic items (new species etc.) 

New snecies ete. can be accepted for publication in .the herbarium when a sufficient 
amLSXwe material (on host or on artificial medium) is rendered and recognized by 
the annointed specialists taking care of the fungus group in question. 

nhv?omlv th? realization of a herbarium as outlined above would be a very large 
underttowLh undoubtedly would have to be built up gradual^. The co-operation of 
mvmiodcaf tlxonomists must be based primarily on vo untary effort The result might 
hi therefore, that some groups would be predominantly represented berause of 
dmint co-operation of some of the appointed specialists than of others But with time 
S can be regulated. It would be the object of the enterprise to make the exsiccatae so 
valuable and iell documented that all the main institutions of the world performmg 
SoloScal and plant pathological investigations would be willing to subscribe for a 
SSpkte set, thus facilitating identical determination of at least the most common and 
the most important plant pathogens. 

{Accepted for publication 30 June 1947 ) 

[ 349 ] 



i4~2o September 1946 

It was most fitting that the first post-war Annual Autumn Foray, the first since 1938, 
should be held in the Jubilee year of the Society; appropriate too that the Yorkshire 
Naturalists’ Union should join forces for the occasion with its younger sister society. 
Headquarters were at the Bungalow Hotel, Sandsend, near Whitby, and, although 
accommodation was a little cramped, everything possible was done for the convenience 
of the party, including the turning of one of the two lounges into a workroom. Some 
eighty members belonging to one or both of the two societies were present, a gratifying 
feature being the number of young enthusiasts who were evidently enjoying their first 
experience of a full-scale foray. 

Conditions were somewhat unusual owing to the abnormal amount of rain which had 
fallen during the summer. Much of the ground was almost or quite waterlogged, so that 
the larger fungi were scarce in most of the areas visited; the fact that these organisms 
require air as well as moisture was only too well demonstrated. Those who arrived late 
on Saturday were soon informed, by members who had reached Sandsend earlier in the 
day and had already carried out a preliminary survey, that Mulgrave Woods were likely 
to be disappointing. The Woods, which lie close to the Hotel, were visited in force on 
Sunday morning, and it was soon evident that the ground, even in the higher portions 
of the area, were almost completely sodden. There was a fine patch of Psathyrella dissemi” 
mta at the entrance to the Woods, and a crop of unusually large specimens of Auricularia 
auricula'judae was found on a fallen branch of Sambucus, but although the total number of 
agarics collected, in this and subsequent searches, was not inconsiderable, most species 
were found as isolated specimens and many were somewhat atypical. 

On Sunday afternoon a large party went to Skelder Plantation, which lies on higher 
ground. In general, fungi were more abundant here than in Mulgrave Woods, but there 
was little variety and nothing of special importance. In the evening Dr Grainger gave 
a paper, illustrated by lantern slides, entitled ‘From Foray to Farm Practice’, giving the 
results of an ecological study of Erysiphe graminis, and showing how observations made at 
forays may be used to help the agriculturalist to combat disease. 

Forge Valley was visited by motor-coach on Monday. A few well-drained areas on the 
steep slopes of the enclosing hills yielded many specimens of Lactarius delidosus and 
Hygrophorm virgineus^ but otherwise finds were few and far between. A high spot of the 
outing was undoubtedly to drive back over the moors in the late afternoon. In the evening 
Mr A. A. Pearson gave a public lecture on ‘ Mushrooms and Toadstools with coloured 
lantern slides, in the schoolroom at the neighbouring village of Lythe, and a goodly 
contingent from the Hotel climbed the mile-long hill to help swell the audience. 

On Tuesday there was another coach trip to Egton, Amcliffe Woods, and Guisborough 
Moor — at least that was the route as planned. The first part of the day’s outing went 
according to schedule and, in spite of some rain, collecting was fairly successful. However, 
when starting on the last stage of the journey, the driver of one of the two coaches, in 
attempting a delayed gear-change, broke a shaft. Fortunately habitation was not far 
a%vay and the stranded half of the party, including the President, whiled away a pleasant 
hour over tea, until a relief coach could be sent from Whitby. The following day there 
was no organized excursion. Many members took the opportunity to see something of the 
ancient port of Whitby, whilst others continued collecting in the Mulgrave area. After 
dinner, the workroom in the hotel was crowded for an ijoformal discussion, organized by 
Br Grainger, on the ecology of fungi, the general conclusion reached being that much 
more work on the subject is required before we can even begin to generalize. 

Thursday brought showery weather and only two members set off for Grinkle Park, 
one of the areas originally planned for a foray. They returned triumphant with repre- 
sentatives of close on 100 species, including a fine selection of brilliantly coloured species 

350 Transactions British Mycological Society 

of Hygrophorus, good specimens— -the only ones collected during the Foray — of Folyporus 
picipes, and the somewhat uncommon Clavaria luteo-alba. 

On the whole the most successful hunting was done by the micro-fungi contingent, as 
there was no lack of interesting material. There were few times in the day, except meal- 
times, without the sound of choppers and saws, and the devotion of certain members to 
their microscopes during ail the hours when out-door foraying could not be indulged in 
was exemplary. In all seventy Pyrenomycetes and fifty-one Hyphomycetes were collected 
and identified, including three new records for Yorkshire. Chaetosphaeria cupuUfera (B. & 
Br.) Sacc., was found in association with Catemlaria cuneiformis [Kichon) Mason on wood 
of Quercus; Rhamphoria tympanidispora Rehm, especially interesting because the ascospores 
bud inside the ascus, was collected on rotten wood of Quercus; and Gomtobotryum fuscum 
Sacc. was found growing on the end of an oak log intended for the fire. The latter fungus 
has been found only three times in this country, the first British record being from an oak 
log in a hotel wood-pile at Haslemere in 1945 {Trans, Brit. myc. Soc. 1946, xxix, 97). 

A full list of the fungi collected during the Foray has been published by Dr Grainger, 
Miss Grainger and Mr W. G. Bramley in the Naturalist (1947, April-June, pp. 83-90). 
The list includes many species which are not to be found in Mason and Grainger’s 
Catalogue of Yorkshire Fungi. 



Four one-day forays were held in the London area. At the first of these, on 28 September, 
Mad Bess Wood, Ruislip was visited, and the impression on entering the wood, noted 
particularly by those who had experienced the scarcity of agarics at the Sandsend Foray, 
was of a welcome abundance of fungi. However, it was soon evident that it was a case 
of quantity rather than variety. Species of Russula, and the small brown species of 
Lactarius, particularly L. quietus and L. subdulcis, were everywhere. It is noteworthy that not 
a single specimen of the usually common Paxillus involutus was found, and only a few 
undersized specimens of Bokttis were collected. Species of Clavaria were unusually plentiful, 
including C. fusiformis, c. cinerea, c. cristata, c. inequalis, c. kunzei, c. rugosa, and c. umhrimlla. 

A joint foray with the London Natural History Society, Ecoiogica! Section, was held 
at Bookham Common on 29 September. South-east and Central Woods were worked in 
the morning and afternoon respectively and a fairly large number of specimens were 
collected, although nowhere was there abundance. Again Paxillus imolutus was missing 
and of Boletus only a few specimens of B. scaber were found. The most interesting discovery 
was on an old and decrepit Pyrus malus. One of the two main stems bore hundreds of caps 
of Marasmius ramealis, although the tree, in spite of its poor condition, was still bearing a 
fair crop of fruit. One patch of abnormally large specimens of Entoloma lividum was found 
under an oak, and several dense clusters of Leotia lubrica were seen. 

On 5 October, Harrow Weald Common and Grim’s Dyke were visited in conjunction 
with the Herts Natural History Society. Amongst the more interesting finds were 
Geaster fornicatus, Amanita porphyria. Boletus custaneus and Peziza aurantia. The Society is 
much indebted to Mr Swallow, of Harrow Weald Grammar School, who assisted in 
making the arrangements, entertained the party to tea in the school, and threw open the 
biological laboratory for identification and display of specimens. 

The season ended with a foray at Oxshott on 12 October. A fairly large number of 
fungi were collected but nothing of outstanding interest was recorded. 

G, Smith 

■' Sec. Foray Committee 

f 351 ] 


27 June 1947 

The Twenty-first Annual Field Day was held at the School of Agriculture, University 
College, Nottingham, Sutton Bonington, by permission of Prof. H. G. Robinson. Nearly 

fifty members and friends attended. 

The programme began at 11.30 a.m. with a demonstration of the soft-fruit plots used 
in training Inspectors for the Ministry of Agriculture’s Certification Schemes. Mr Martyr 
then conducted the party on an interesting tour of the Horticultural Department during 
which there was a lively discussion on some of the diseases present. 

The members were the guests of the School of Agriculture at lunch. After lunch the 
President of the Society, Prof. C. G. G. Chesters, thanked Prof. Robinson and his staff 
for their hospitality and for arranging a most interesting programme. In replying, 
Prof. Robinson outlined the developments which were taking place at the School of 
Agriculture. In the afternoon Dr A. R. Wilson gave an account, illustrated by exhibits, 
of methods for storing potatoes which he had seen in the United States. He also discussed 
the problems of potato storage under English conditions and the work that being 
done on them. 

Under the expert guidance of Mrs N. McDermott a tour was also made of the potato 
plots. This proved of great interest, although it was somewhat curtailed by a thunder- 
storm. After tea in the Canteen most members returned to the potato plots where 
Dr Gregorys demonstrated the results so far obtained on the spread of potato virus diseases. 
Members also took the opportunity of seeing other potato plots in which they were 
particularly interested. The party dispersed at about 5.30 p.m. 

H. E. Croxall 
Sec* Plant Pathology Committee 


J^orth American Species of Mycena. By Alexander H. Smith. University of 
Michigan Studies, Scientific Series, vol. xvn. (Ann Arbor, Michigan: 
University of Michigan Press; London: Oxford University Press, 
1947.) xviii-l-521 pp., 56 text-figures, 99 uncoloured plates. $6.00. 

With the increased use of the microscope in the study of the higher fungi ideas as to their 
classification have in recent years undergone a revolution, but are still in a state of flux. 
Many new genera have been established, while, on the other hand, some old genera have 
been united on the basis of microscopic characters, as, for instance, the genus BJwdophyllus, 
which includes all the pink-spored Agarics with angular or nodulose spores. One of the 
earliest genera of Agarics to be studied microscopically was Mycena, for many species of 
which Schroeter in 1889 described cystidia and spores. Since that time Mycena^ has been 
the subject of several revisions. Von Hohnel, in 1913, proposed a classification of the 
European species based on the characters of the cystidia, and Lange (19^4 ^ 93 ^) 

divided the section Eumycena into two groupe, Ciliatae and Granulatae, according as the 
cystidia were smooth and more or less pointed or rounded, with spines or finger-like 

352 Transactions British Mycological Society 

appendages, Oort, in 1928, published a revision of the Mycenas of the Netherlands, and 
Kiihner in 1938 produced an elaborate monograph of the European species of Mycena^ 
in which he emphasized the importance of not only cystidia and spores but also the 
structure of flesh and gills, the covering of the pileus and stem, and the iodine reactions 
of both spores and hyphae. In America Kauffman, in 1918, described cystidia and spores 
for the species of Mycena found in Michigan, and Beardslee and Coker, in 1924, gave an 
account of the Mycenas of North Carolina with drawings of microscopic detail. Between 
1935 ^tid 1939 A, H. Smith published some preliminary studies of North American 
Mycenas, and this author has now completed a monograph of the North American 
species of the genus. 

As might be expected, the North American flora is much richer in species than is the 
European; the present work includes descriptions of 218 species, as compared with 143 
recognized by Kiihner for Europe. Most of the European species are found also in 
North America, but the student who is using this book, along with say Kuhner^s mono- 
graph, may find differences of interpretation of some of the old species. Unfortunately, 
the older mycologists seldom thought of preserving specimens of the Agarics which they 
described, and the descriptions alone are often insufficient to indicate precisely which 
of two or more outwardly similar species was in question. Thus it comes about that 
different authors may interpret some of Fries’s species in different senses. The species 
which is known in this country, and to Lange, Oort and Konrad and Maublanc as 
Mycena filopes is described by Kiihner under the name of M. mtilis Fr., while Kuhner’s 
M. filopes is what these other authors call M. vitilis. Smith, in the present work, distin- 
guishes a species with no particular odour which he calls M. filopes from one with a distinct 
smell of iodoform which is referred to M. iodxolens LundelL Again, a student wffio is trying 
to identify the common Mycena of pine woods with red-edged gills and a distinct nitrous 
smell would, if using either Kiihner or Smith, come to the name M. capillaripes Peck. 
Yet in this country this species is usually referred to M, rubrormrginata Fr., a name which 
is given to a different fungus by the authors mentioned. One of the most urgent needs 
for mycological nomenclature is some guidance as to the selection of type, or the method 
of fixing the application of a name, in such cases as these wffiere no authentic or type 
specimen exists. If no agreement can be reached, it would be better to discard some of 
these old names, as is suggested by Smith for * Mycena cMorantha Fr,’ It is a relief to find 
that Dr Smith is, on the whole, conservative both in his conception of the genus and in 
his interpretation of old species. For purely American species his task has been easier, 
since the type specimens of many of these exist and he has been able to examine them. 

After an exposition of his broad concept of the genus Mycena, wffiich includes certain 
species formerly described as Collybia or Omphalia, the author discusses in detail the various 
diagnostic characters used for species and describes his technique. Then follows an 
explanation of his grouping of the species into four subgenera, and an attempt is made to 
place related species together in small groups or stirpes within the adopted subgenera 
and sections. 

The descriptive portion of the work, which occupies pages 43-452, is provided with 
keys to subgenera, sections and species, and includes full descriptions of all species known 
to the author. He has also added (pages 453-470) emended descriptions of MurrilFs 
tropical and subtropical species of Mycena and OmphaUaj drawm up from examination of 
the type specimens. 

Details of microscopic structure are shown in line drawings, and at the end of the book 
are 98 plates depicting in half-tone reproductions from photographs the habit of fresh 
specimens of those species known to the author at first hand. < 3 ne would have preferred 
coloured figures, but in these days the expense of colour printing is a limiting factor. 
With the help of the photograjphs and the microscopic characters together it should be 
possible to determine any given species with some degree of certainty. The book is 
completed by a list of doubtful or excluded species and a bibliography of i2|- pages. 
As in other books published by the University of Michigan Press the format is good, the 
text and drawings clear and easy to consult. The serious student of the genus will find 
the work valuable' and interesting to compare with Lange and Kiihner. Especially 
useful are the critical observations which follow the description of each species. 

E. M. W. 

[ 353 1 



Joint meeting with the Genetical Society held in the rooms of the T.innean Society of 

London on Saturday, 8 March 1947. 

S. Dickinson (School of Agriculture, Cambridge). Fungal Genetics. 

The study of fungal genetics has hitherto been a study of haploid inheritance. The size 
of the nuclei has prevented any considerable advance by joint cytological and genetical 
studies. The association and dissociation of nuclei in multinucleate types has been a 
fruitful subject for investigation. The linear and naked arrangement of the products of 
segregation in such fungi as the Smuts may provide a profitable line of research on polarity 
or biased segregation. 

Elizabeth Blackwell (Royal Holloway College). The morphology of the 
strains of certain heterothallic fungi and the terminology of the organs 
of fusion. 

A review of the many different mechanisms which have been evolved in the fungi for 
bringing haploid nuclei together, and ultimately forming the zygote, suggests a critical 
consideration of the terms in use for the organs of syngamy: names of organs found in 
algae and assuming homology with them. The unit of the fungus is a hypha, not a cell, 
and liyphae readily fuse with one another. 

A few examples taken from the ‘higher fungi’ will serve. In Pyromma conjluens and 
Ascobolus magnijicus there are specialized hyphae of fusion called, unfortunately, ‘antheri- 
dium’ and ‘ascogonium’, sometimes even ‘oogonium’. In the former (homothallic) 
species these arise close together: in the latter (heterothallic or better, homothallic and 
self-sterile) species they develop only on contact of the strains though both strains bear 
both types of hypha. In the former there is no risk of their not meeting and it is significant 
that here there are no accessory spores. The strains of the latter species are widely 
dispersed by papulospores, and so brought together. In Pleurage {^Podospora) anserina 
there is only specialized hypha, the so-called ‘trichogyne’; and the haploid nucleus of 
opposite strain comes direct to it in a microconidium, transported by air, or water, or 
insects, and is absorbed by it. This is a veiy efficient mechanism. Fusion of conidium 
with conidium occurs in yeasts and and this association of strains by two trans- 

portable units is even more efficient. In species of Puccinia there is ready fusion between 
unspecialized hyphae, thus bringing haploid nuclei into the same segment; but in hetero- 
thallic species, e.g, P. graminis and P. helianthiy the chances of association of different 
strains are increased by the production of a transportable unit, viz.: the insect-borne 
pycnidiospore, which is absorbed by a flexuous hypha grovdng out from the pycnidium 
of opposite strain. This is equivalent to ‘spermatium’ and ""iTichogYnt^ oi Pleurage anserina^ 
but there is no need to call these ‘ sex organs Indeed the colour, scent and nectar of the 
pycnidium recall pollination, the transport of a spore, rather thnn fertilization, the fusion 
of ovum and sperm. It is significant that the homothnllic Puccinia malvacearum does not 
produce pycnidia. In the heterothallic Coprinus lagopus there is ready fusion of hyphae, 
and again the conidia produced on the haploid mycelia increase the chances of strains 
meeting. Three phenomena in these ‘higher fungi’ deserve our attention: (a) the delay 
of fusion of associated haploid nuclei; {b) the development of a hook hypha: the ‘crozier’ 
of ascomycetes, the ‘clamp connexion’ of basidiomycetes; (^) the fact that, except in the 
immediately after fusion, meiosis follows: there is no resting spore with a diploid 

It would seem that in this unique group of the fungi, quite original and dependent 
methods of breeding and cross-breeding are found, analogous to, but not homologous 

354 Transactions British Mycological Society 

with, the methods of plants in general. In conclusion, {a) the different so-called 'sex 
organs ’ may have been derived independently from vegetative hyphae which had already 
developed a tendency to fuse with one another; {b) the conidium (oidium, papulospore, 
spermatium, etc.) is efficient in effecting the wide distribution of a haploid strain, even 
when it cannot germinate of itself h\xt must needs be absorbed by a receptive hypha, and that 
this is of importance not only in the ultimate association of different strains, but in 
increasing the chances of hybridization; {c) for the mere association of haploid nuclei 
neither heterothallism nor the production of special organs of fusion is necessary. 

C. Robinow (Strangeways Laboratory, Cambridge). On the staining of 
nuclei in bacteria and fungi. 

In preparations stained by conventional methods the nuclei of bacteria are usually 
invisible. The reason for this lies in the strong affinity of the cytoplasm for basic stains 
which is due to its high content in ribonucleotides. Extraction with normal hydrochloric 
acid at 57-60° G. for 5-10 min. removes the ribonucleotides from the cells (Vendreiy, 
1946) but does not lower the affinity for basic dyes of the nuclear structure. If the ceils 
are stained after hydrochloric acid extraction, little of the stain is retained by the cytoplasm 
and the nuclei stand out very clearly. Of many basic dyes tried Giemsa’s stain has proved 
most satisfactory; the effect of the stain parallels exactly, but is more brilliant than that 
of the Feulgen test. Giemsa-stained nuclear structures in hydrochloric acid extracted 
material appear slightly larger than in Feulgen preparations. 

The same result has recently been obtained by digesting fixed bacteria with the 
enzyme ribonuclease (Tulasne, Univ. Strasbourg) . 

In bacterial spores it is not a basophilic cytoplasm which prevents a view of the interior 
but the impenetrable spore membrane (of unknown composition). The staining properties 
of bacterial spores are radically altered after 10-20 min. in normal hydrochloiic acid at 
room temperature. This simple expedient renders the spores transparent and reveals their 
relatively large refractile nuclei. The nuclei are superficially attached to, not contained 
in, the much less refractile, dormant cytoplasm. The chemical basis of the effect of 
hydrochloric acid on the spore membrane is not known. Changes in the arrangement of 
the nuclear structures precede the formation of spores. The processes an example of 'free 
ceil formation’ (in the cytoplasm of the spore-forming bacterium). 

In the study of the nuclei of yeast cells extraction with warm hydrochloric acid before 
staining has proved quite as useful as in bacteria and has allowed the staining of the 
chromatinic bodies first described as nuclei by Badian, 1937. Schizosaccharomycetes 
have been found more rewarding subjects than Saccharomyces cerevisiae. 

Hydrochloric acid extraction greatly facilitates the staining of the nuclei in whole- 
mount preparations (on cellophane) of fungal mycelia. 

The method was illustrated with photomicrographs of bacteria, yeasts, and hyphae 
and conidia of Penicilliim notatiim, Oospora lactis, and Mucor kiemalis. 

A. F. Parker Rhodes (Department of Genetics, Cambridge). Pheno- 
genesis in fungi. 

The action of genes in producing the visible characters of an organism is called Piieno- 
genesis; its study belongs partly to biochemistry and partly to embryology. This paper 
considers the theory of the process as applied to fungi. The theory is based on two concepts, 
both of biochemical origin, viz. : Autonomy and Biodynamic Field. By their help much 
of the experimental observations on the development of plants and animals can be 
reduced to coherence. 

Fungi differ from both animals and plants. Many groups of fungi are built of hyphae, 
and their tissues are thus usually anisotropic ; this means tiiat unlike other oi'ganisms they 
must have fields of two kinds, synhyphic (gradients along preferred direction of hyphae) 
and anthyphic (gradients across). Because inter-hyphal diffusion is usually easy ant- 
hyphic fields are open to environmental interference, whereas s>mh>jphic ones can be 
made autonomous. ■ It' can, be shown that. such fungi can tolerate a much higher mutation 
rate among those genes which are called into play by synhyphic fields than among those , 

Proceedings 355 

calied into play by anthyphic ones; whence characters governed by anthyphic fields are 
either environmentally variable or of considerable stability. 

Evidence is presented that single-gene variants in nature nearly always concern 
synhyphal characters. Genetic evolutionary implications are mentioned, and the need 
for experimental research stressed. 

G. PoNTEGORVO (Department of Genetics, University of Glasgow). 
Genetical aspects of heterokaryosis. 

Heterokaryosis is the coexistence of nuclei not all genetically alike in a syncitial tissue. 
It may occur in any syncitial tissue of higher organisms, as well as of colonial micro- 
organisms, as a consequence of mutation. In most filamentous fungi, however, hetero- 
karyosis is brought about regularly through an appropriate mechanism. Anastomoses 
between hypiiae occur readily and if hyphae that fuse differ in the kind of nuclei they 
carried, a hypha with nuclei of two, or more, kinds may result. Heterokaryosis is wide- 
spread in homothailic, hetez'Othailic and ‘imperfect’ species or strains. 

Since the nuclei of different kinds in a heterokaryotic hypha can segregate (e.g. at 
‘cell’ division, or in the formation of conidia, etc.), the alternation of hyphal fusions with 
segregation of whole nuclei provides a mechanism of genetical recombination additional 
to that of alternation of meiosis and fertilization known from classical genetics. The two 
mechanisms may or may not coexist in any particular species or strain, and the genetics 
of natural populations in fungi must therefore consider genetic systems of the classical 
types (based on sexual reproduction alone) as well as genetic systems based on hetero- 
karyosis alone (or almost so) and mixed systems in which both heterokaryosis and sexual 
reproduction play important parts. 

The physiological genetics of heterokaryons presents many new problems, e.g. (i) the 
action of allelomorphs carried in different nuclei in the same hypha (dominance, localized 
action, etc.) ; and (2) the mechanisms of control of the ratios of diferent kinds of nuclei 
within a hypha by internal and external conditions. Examples were given of how these 
and other problems are investigated by means of artificially synthesized heterokaryons 
carrying two or more kinds of ‘labelled’ nuclei. 

K. Mather (TheJohnlnnesHorticulturallnstitution^London). Genetical 
flexibility and the breeding system in fungi. 

The genetical flexibility of a sexually reproducing species depends upon the bringing 
together, by recombination, into one individual of genes from different parents. Such 
recombination can occur only in heterozygotes, resulting from cross-breeding. Flexibility 
is thus greatest in regularly cross-breeding species and least in regular in-breeders. Genes 
coiitroliing the type of mating have an adaptive value through this effect on flexibility. 

Gross-breeding is stimulated at two levels by genes in fungi. The commonest kind of 
heterothally prevents self-mating at the haploid level, where inbreeding would lead to 
immediate and complete loss of flexibility through homozygosis. Genetically more 
complex types of heterothally also reduce the relative frequency of mating between 
hapioids from the same diploid zygote, so reducing inbreeding at the diploid level, where 
the loss of flexibility is slower but eventually complete. 

Inbreeding can be advantageous by increasing the frequency of a highly adapted type, 
at the expense of flexibility. Inbreeding mechanisms appear to have been superimposed 
on relic cross-breeding mechanisms in fungi as in Angiosperms. 

Some iuiigi have abandoned the sexual cycle and secure by heterokaryosis a kind of 
flexibility which would be impossible in highly differentiated organisms. The sexual cycle 
and its associated outbreeding devices, may persist in species which have adopted 

H. L. K. Whitehouse (Botany School, Cambridge). Evolutionary aspects 

of heterothaliism and heterokaryosis. 

The simplest type of heterothaliism, controlled by a gene locus with two alleloniorphs, 
is found in the Mucorales, in at least seven orders of Ascomycetes, and in the Uredmales 


356 Transactions British Mycological Society 

and Ustilaginaies. Heterothallism controlled by multiple allelomorphs at one or two loci 
has been recorded only in the Hymenomycetes (including the Tremellales), but probably 
also occurs in the Gasteromycetes*. Published data indicate that of approximately 130 
species of -Hymenomycetes that have been tested, 50 % have two loci for heterothallism, 
40% one locus and 10% are homothallic, with no significant differences between 
different families. That Auricularia and Exidia in the Tremellales have multiple-allelo- 
morph heterothallism (Barnett, 1937) like the Agaricales, implies that the Tremellales 
probably have a much closer affinity with the remainder of the Hymenomycetes than with 
the Uredinales with their two-allelomorph heterothallism. 

In some heterothallic fungi, such as Neurospora crassa, heterokaryosis appears to be 
restricted to between strains of one mating t^e (Sansome, 1946). In many others, 
association of nuclei of different mating types is brought about at an early stage of the 
life history by the fusion of vegetative hyphae, and in these cases it would appear that 
differentiated sex organs, if present, must be inessential for the completion of the life 
cycle, e.g. Uredinales (Brown, 1935)* Using the term heterokaryosis in the broad sense 
to cover all cases of hyphae containing two or more genetically unlike nuclei, irrespective 
of whether the association has any ultimate sexual significance or not, then in the selective 
advantages to a species of unrestricted heterokaryosis can be seen a possible explanation 
for the apparent progressive loss, first of the sex organs as structures essential for sexual 
reproduction, and then of the organs themselves, as one proceeds from the lowest to the 
highest groups of fungi. 

In those heterothallic species in which heterokar^^osis is restricted to mycelia of one 
mating type, multiple-allelomorph heterothallism would be disadvantageous compared 
with the two-allelomorph type in so far as it would restrict the possibility of heterokaryosis 
to a narrower field. Thus it is improbable that multiple-allelomorph heterothallism 
would acquire a selective advantage in evolution until the condition of unrestricted 
heterokaryosis had first been evolved. 


Barnett, H. L. (1937). Mycologia^ xxix, 626-49. 

Brown, A. M. (1935). Phytopathology, xkv, 1085-90. 

Sansome, E. R. (1946). BulL Torrey BoL Ci. Lxxm, 397-409. 

* JSfote added in proof. Fries (1940) found multiple-allelomorph heterothallism in the 

Fries, N. (1940). Symbolae Bot, Upsal, iv, 1-39. 


A visit was made to the Brewery and, by kind invitation of Mr H. J. Bunker, the Brewery 
Research Department of Barclay Perkins & Go. Ltd. of Southwark on the afternoon of 
Friday, 18 April 1947. 

Meeting held in the Department of Botany, University College, Nottingham, at 1 1 a.m., 
on Saturday, 28 June 1947. The President, Prof. C. G. C. Chesters, in the Chair. 

Morning Session 

Mrs E. Sansome. Mutation in Fimgi. imperfecti with special reference to^ ^ Penmiiium 

G.'C.'M."' Harris.: The mo^rphology of PenicUHum chrysogenum^ in submerged culture.. 

Afternoon Session 

P, T. Thomas. Some aspects of the cytology of Fungi. 

F. L. Drayton and J. W. Groves, A new heterothallic Stromaiima, 

F. C. Atkins. A new species of Vertkillium on mushroom in England* 

Proceedings 35 j 

Meeting held in the Department of Biology, Chelsea Polytechnic, London, at ii a.m., 
on Saturday, 25 October 1947. Mr G. Smith, Vice-President, in the Chair. 

The meeting took the form of an Exhibit from the Herbarium of the Imperial Myco- 
logical Institute, Kew, of some classical British moulds. The exhibit, which had been 
arranged by S. J. Hughes and M. B. Ellis, was introduced by E. W. Mason. H. A. Dade 
showed specimens from the Institute’s culture collection and Miss J. Hickman demon- 
strated the indexing system used in the herbarium. 


The fifty-first Annual General Meeting was held in the rooms of the Linnean Society 
of London at 12 noon on Saturday, 6 December 1947, President, Prof. G. G. G. 

Chesters, in the Chair. 

Mter the Minutes of the previous Annual Meeting had been read and signed the 
President recorded with deep regret the deaths of two members, Mr G. H. Murray, who 
died when a prisoner of war in 1942, and Mr J. Wilton; and reviewed the year’s activities. 

The Treasurer, Mr W. Buddin, then presented his annual statement and drew atten- 
tion to the increasing cost of the Transactions^ the price of which to non-members had 
had to be increased. He urged contributors to present their results concisely and to make 
every effort to reduce the cost of corrections to the proofs which had been excessive for 
recent numbers. 

The following Officers and new Members of the Council for 1948 were then elected: 
President^ A. E. Muskett; Vice-Presidents, P. O’Connor, and the two past-Presidents, 
C. G. C. Chesters and J. Ramsbottom; Secretary, C. J. Hickman; Foray Secretary, G. G. 
Ainsworth; Treasurer, W. Buddin; Editors, B. Barnes and W. C. Moore; Members of the 
Council, T, E. T. Bond, Miss M. P. English, G. Smith, and E. R. Wallace (to replace 
R. W. G. Dennis, C. G. Dobbs, C. J. Hickman, and G. Samuel). A. S. Boughey, R. W. 
Marsh and Miss F. J. Moore were elected as new members of the Plant Pathology 

On a motion from the Chair, E. W. Swanton was elected as an Honorary Member. 
Nine new Members and three new Associates were elected, making fifty new Members 
and twenty new Associates for the year. 

A recommendation to the Annual General Meeting from an ordinary meeting of the 
Society, held during the Autumn Foray at Exeter, that the presidential address should 
in future be delivered at the Autumn Foray and that the Annual General Meeting should 
be held later in the year was discussed. It was agreed that from 1948 the presidential 
address should be given at the Autumn Foray unless the President should decide other- 

After the programme for 1948 had been discussed, the meeting adjourned until 
2 o’clock when the President delivered his address, ‘ Concerning fungi inhabiting the 
soil’; a vote of thanks to Prof. Chesters concluded the meeting. 

G. C. Ainsworth, Secretary 

for the year ending 30 June 1947 

1946 £ s. d. £ s. d, 1946 

I July Balance brought forward ... ... 925 16 3 16 July Wreath for Carleton Rea, Esq. .. 

26 July Grant from Royal Society 75 o o 14 Aug. Purchase of 2!% Defence Bonds .. 


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[ 359 ] 


New names are indicated by an asterisk 

Abies sibiricae, 235 
Absidia giauca Hagem, 247 
Acer pseudoplatanus (sycamore), 288 
Achlya Nees, 257, 262, 264 
polyandra Hiideb., 263 
Acocephalus nervosus, 299 
Acremonium Link ex Fr., 302 
album Preuss, 302 
araneariim Fetch, 292, 302 
cieoiii Wize, 298 
tenuipes Fetch, 292 
Acrophyton tuberculatum Lebert, 291 
Acgeria pyri, 297 
Aegerita Fers. ex Fr., 143 
insectorum Fetch, 298 
Aegopodiuin, 143 
Agarics, British, index to, 147 
Agaricus (Omphalia) affricatus B. & Br., 201 
agathosmus Fr., 196 

(Omphalia) alutaceus Cooke & Massee, 201 
(Goilybia) andersoni, 199 
atomatus, 207 
babingtonii Blox., 206 
(Entoioma) bulbigenus B. & Br., 205 
crassifolius Berk., 197, 198, 208 
fumosus Fers,, 197, 198 
(Coliybia) leucophaeus B. & Br., 21 
(Goilybia) muitijugus B. & Br,, 20 
(Coliybia) nepheiodes B. & Br., 19 
(Mycena) pailido-rubens B. & Br., 24 
pachyphyllus, 198 
paupercuius Berk., 199 
(Entoioma) persooni Du Port, 205 
pei'soonianus Phill., 205, 208 
pustuiatiis Fers., 196 
sericeus Fers., 205 
stevensonii B. & Br., 199 
Agriotes,. 298 
sptitator, 289 
Agrostis tenuis, 277 
Agrotis ripae, 294 
segetum, 301 

Akauthomyces aculeata Lebert, 296 
sphiiigum (Schw.) Fetch, 291, 298 
Aleurodiscus Rabenh., 241 
Aleyrodes vaporarium, 293 
Allium cepa (onion), 341 
Alnus (alder), 291 
Alternaria Nees ex Wallr., 282, 346 
solani (Ell. & Martin) Sor., 341 
Arnani ta muscaria (L.) Fr., 1 08 
porphyria (i4ib. & Schw.) Fi'., 350 
Ambrosia fungus, 13, 14 
Amelanchier, 89 
Amomuni, 38 

Androsaceus (Pers.) Pat., 243 
Aneura, 7, 8, 9 
Angelica, 143 

Annual General Meeting, 1947, 357 
Anthoceros, 8 

Antifungal substances from moulds. 11. The 
effect of patulin on the growth of various 
plant parasites, by G. Smith, 136 
Antirrhinum majus, 86 
Aphrophora alni, 299 
spumaria, 288 

Araeopus pulchellus, 297, 304 
Armillaria jasonis Cooke & Massee, 191, 

Arthrinium sporophlaeum Fr., 143 
Aschersonia mellea B. & B., 38 
Ascobolus magnificus Dodge, 353 
Ascochyta Lib., 346 

bohemica Kab. & Bub., 89 
cinerariae Tassi, 89 
dahliicola (Brun.) Petr., 90 
impatientis Bres., 90 
(Diplodina) lycopersici Brun., 89 
weissiana Allesch., 90 
Asparagus, 286 
Aspergillus Mich, ex Fr., 30 
depauperatus Fetch, 295 
Asterina congesta Cooke, i 
Asterodon, a clue to the morphology of fungus 
fruit-bodies: with notes on Asterostroma 
and Asterostromella, by E. J. H. Corner, 


Asterodon Pat., 234, 241, 244 
ferruginosus Pat., 234, 235-9, 241 
Asterostroma Massee, 235, 241, 242 
degenerans Bres., 241 
laxum Bres., 241 

muscicolum (B. & C.) Massee, 241 
ochroleucum Bres., 241 
Asterostromella v. Hoehn. & Litsch., 241, 243, 

Athysamis gnsescens, 288 
Atkins, F. G., *A Verticillium disease of cultivated 
mushrooms new to Great Britain’, 126 
‘A new species of Verticillium on mushroom 
in England’, 356 

Auricularia auricula-judae (L, ex Fr.) Schroet., 
349 , 

Autumn forays, 1946, 349 
Avena (oats), 59, 341 

Beaumont, A., ‘The dependence on the 
weather of the dates of outbreak of 
potato blight epidemics’, 45 
Beauveria bassiana (Bals.) Vuill., 294, 299 
densa (Liixk) Picard, 294, 299, 302 
effusa (Beauv.) Vuill, 294 
Bertia moriformis (Fr.) de Not., 143 
Bicricium naso Sorokin, 84 
transversum Sorokin, 84 
big bud, 293 



Blackwell, E.j ‘The morphology of the strains 
of certain heterothallic fungi and the 
terminology of the organs of fusion’, 353 
Blepharoptera serrata, i298, 299 
Boletus Dill, ex Fr., 108, 350 
castaneus (Bull.) Fr., 350 
scaber (Bull.) Fr., 350 
Bombardia iunata, 77 
Bombus pratorum, 294 
Botryosphaeria pulicaris, f. baccata, 87 
f. mori albae, 87 

Botryosporium iongibrachiatum (Oudem.) 
Maire, 90 

Botryotrichum Sacc. & March., 282 
Botrytis Pers. ex Fr., 116, 117, 119, 247, 346 
allii Munn, 247 

cinerea Pers. ex Fr., 117, 136, 137, 139, 340, 


eriophyes Massee, 293 
fabae Sardina, 340 
Brassica, 286, 290, 300 
Bremia Regel, 116, 119, 123, 124, 125 
lactucae Regel, 1 12, 1 14, 1 15, 1 16 
British Agaricaceae. Little known species. 
I. Leucosporae and Rhodosporae, by 
R. W. G. Dennis, 191 

British Agarics and Boleti; Revised list of, 
by A. A. Pearson and R. W. G. Demiis, 


British entomogenous fungi: Revised list of, by 
T. Fetch, 286 

British fungi, by W. C. Moore, 86 
British fungi parasitic on Lichens, List of, by 
W. Watson, 305 

British Plant diseases, List of, 340 
Bromus inermis, 249 
Brownea grandiceps, 14 
Bryophytes, fungal diseases of in Mysore, by 
M. J. Thirumalachar, 7 

Calamus, 38 
Calceolaria, 137 
Calliphora vomitoria, 289 
Calonectria graminicola Wollenw., 341 
nivaiis Schaffnit, 341 
Caltha palustris, 143 

Calyculosphaeria tristis (Fuckel) Fitzp., 143 
Camarophyllus (Fr.) Karst., 197 
Camellia, 88 
sinenesis (tea), 17, 18 
Campanella cucuilata (Jungh.) Lloyd, 36 
Campanula betulacfolia, 89 
medium, 89 
ranieri, Bg 

Canter, H., ‘On Myzoiytitim megastonmm De 
Wildeman’, 80 

‘Studies on British Chytrids, II. Some new 
monocentric chytrids’, 94 
‘Studies on British Chytrids. III. 
zidium willei hiywtnmBl znd MhizophMum 
Cantharelius Adans. ex Fr., 39 
capensis, 35, 36 
eiegans B. & Br., 22 
inaequalis B. & Br., 39 
rabidus (rubidus), 26 

Cantharis, 289, 290 
lateralis, 294 

Gapitophorus fragariae, 293 
Cards, perforated, for preliminaiy identifi- 
cation of fungi, by W. P. K. Findlay, 

Garex, 142, 143 
riparia, 143 
Cercospora Fres., 346 
Cercosporelia primulae Allesch., 90 
Cephalosporium Gorda, 291 

(Acrostalagmus) aphidicoia Fetch, 293 
(Acrostalagmus) coccoram Fetch, 293 
(Acrostalagmus) dipterigenum Fetch, 294 
(i\crostalagmus) eriophytis (Massee) Fetch, 


(Acrostalagmus) lefroyi Hoxme, 293 
militare Kobayasi, 291, 294 
(Acrostalagmus) muscarium Fetch, 293 
(Acrostalagmus) subclavatum Fetch, 293 
Ceratostomella pini Muench, 136, 137, 139 
Chaerosphaeria phaeostroma (Dur. & Mont.) 
Fuckel, 143 

Chalaropsis thielavioides Peyron, 341 
Chionaspis, 293 
salicis, 292, 298 

Chytridium A. &'aun, 104^ 128 
characii Scherffel, 1 28 
’^'cocconeidis H. M. Canter, n.sp., 102, 103, 

curvatum Sparrow, 104 
iagenuia A. Braun, 104 
spirotaeniae Scherffel, 128, 129, 130, 


versatile Scherlfel, loi, 102, 104 
versatile Scherffel va,r. acauHs H. M, 
Canter, n.var., loi, 102 
Ciboria Fuckel, 346 
Cicinnobolus ulicis Adam, 86 
Cineraria maritima, 89 
Citrus, I 

Cladium, 142, 290 
mariscus, 143 

Cladochytrium Nowakowski, 7, 8, 9 
*aneurae Thirumalachar, nisp., 8 
nowakowskii Sparrow, 8 
replicatum Karling, 9 
Cladophora, 95 ^ 

Ciadosporium Link ex Fr., 346 
aphidis Thuem., 298 
■herbarum Link, 56, 63, 91, 298 
Clasterosporium caricinum. Schw., 142, 143 
Ciavaria Vaill. ex Fr., 350 
cinerea (Bull.) Fr., 350 
cristata (Holmsk.) BV., 350 
fusiformis (Sow.) Fr., 3,50 , 

. inaequalis (Muller) <iueL, 350 
kunzei Fr., 350 
rugosa (Bull.) Fr., 350'' ' 
setiformis VaM, 299' 
umbrineila Sacc., 3,50 
Clitocybe clavipes (Pers,). Fr., 195, 208,, 
infundibul,iformiS' (Schaeff.) .Fr,, 201 ' 
.obs.curata Cooke, 19.5, .208 
occulta Cooke, T 92 .. 
p.ergamenus Cooke,'' 192 ' 



Clitocybe {continued) 
subiiivolutus W. G. Smith, 195 
zygophila Cooke & Massee, 192 
Glitopiius (Fr.) QueL, 202 
caiicrinus Fr., 206, 208 
pleurotelioides (Kuimer) Josseraud, 202, 208 
.sarnicus Massee, 206, 208 
stilbocephalus B. & Br., 207, 208 
Ciosterium, 130 
costatiim, 80 
rostratuiii, 80 
Ciytocerus oceilaris, 295 
Cocconeis pediculus, 102 
Colletotriciium Gorda, 346 
CoIIybia^(Fr,) QiieL, 19, 197, 205, 352 
eustygia Cooke, 197, 20B 
leucophaea, 30 
muscogena (Schiiin.'*! Fr., 200 
piatyphylia (Pers.) Fr,, 195 
psathyi'oides Cooke, 198, 208 
rancida Fr., 197 
stevensonii B. & Br., 198, 208 
stupparia B. &. G., 29 
thelcphora Cooke & iMassee, 192 
Committee: 'Disease Measurement in Plant 
Pathology’, 343 

Committee: ' Emt^ndations to the third edition 
of the List of Common British Plant 
Diseases’, 340 

Committee : ' Measurement of potato blight ’,140 
Coniophora putcana (Sebum.) Karst., 92 
Coniothecium amentacearu.m Corda, 143 
Coniothyrium Sacc., 350 
Conocephaliiin, 143 
Goprinus (Pers. ex Fr.) S. F. Gray, 143 
lagopus Fr., 353 
Cordyceps (Fr.) Link, 294, 299 
davuiata (Schw.) Ell. Everh., 292 
entomorrhiza (Dicks.) Link, 291 
fourquignoiii QtieL, 292, 296 
fuliginosa Ce$., 297 
gracilis 2 vloat. & Dur., 291, 295 
mcmorabilis Ces., 304 
militaris (L.) Link, 291, 294 , 
pistillariaeformis B. & Br., 292 
sphecocephala (KL) Gtx>ke, 291, 296 
sphingurn (Schw.) B. & G., 291 
swantonii A. L. Smith, 295 
thaxleri Mains, 296 
tubcrculata (Lcbert) Mairc, 291, 296 
Corner, i*i J'. H., ^hkrodon, a clue to the 
morphology of fungus fruit bodies, with 
notes on Asin&siromn and Astmstromdla\ 

234 ^ 

Corticitim (Pers. ex rr,), 143, 23^5 
nivcocrenicum v. Iloelin. & Litsch., 143 
niseocremeum Bres., 143 
solani ^Frill, & Delacr.) Bourd, & Galz., 342 
Cortinarius (Pens, ex Fr.) S. F. Gray, 108 
Corymbites cuprous, 295, 298 
Coiyniebacterium diptheriae, 223 
michiganense (E. F. Sm.) fi L. Jens., 341 
Cosmarium contractum, 84 
CrataegiLs (Hawthorn), 295 
Crepidotus melkus (B, & Br.) Fetch, 38 
Crepis, 1 16 

Croxall, H. E., ‘ The twenty-first annual Plant 
Pathology Field Day. 27 June 1947’ 351 
Cucumis (cucumber), 340 
Gucurbitaria S. F. Gray ex Grev., 346 
Curcuma, 5, 6 

Cylindrocarpon Wollenw., 282, 346 
Gylindrocystis brebissonii, 130 
Cylindrodendrum sufFultum Fetch, 295 
Cylindrophora aranearum Fetch, 290, 292 
Gyphella pulchra B. & Br., 38 
Cyrtoneura estabulans, 292 
Cystopus Lev.. 346 

Dactylis glomerata, 277 
Dahlia variabilis, 90 

Daldinia concentrica (Bolt, ex Fr.) Ces. & 
de Not., 277 

Dasyscypha controversa (Cooke) Rehm, 143 
Dematium hispidulum Fr., 143 
Dendrocalamus giganteus, 33 
Dennis, R. W. G., ‘Some little-known British 
species of Agaricaceae. I. Leucosporae 
and Rhodosporae’, igt 
Diaporthe eres Nits., 143 
Diatrype Fn, 277 

Dickinson, S. ‘Fungal Genetics’, 353 
Didyniella Sacc., 346 
Diplocarpon Wolf, 346 
Diplodina (Ascochyta) fibricola Sacc., 89 
Disease Measurement in Plant Pathology, 343 
Doronicum, 86 

Dovaston, H. F., ‘A new species of Pyrenophora 
from Italian ryegrass’, 249 
Downy mildew disease of the cultivated lettuce, 
by H, Wild, 1 1 2 

Drayton, F. L. & Groves, J. W., ‘A newhetero- 
Stromatinia^ , ’ 

Ecology of Erysiphe graminis DC., by J. 

Grainger, 54 
Elephant hair blight, 34 
Elsinoe Racib., i, 6 

bitancourtiana Thirumalachar, r 
fawcettii Bitan. & Jenkins, i 
Emendations to the tLird edition of the List of 
Common British Plant Diseases, 340 
Empusa Cohn, 298 
acaridis Fetch, 287 
apicuiata Thaxt,, 287 
aulicae Reichardt, 286 
culicis A. Braun, 286 

forficulae (Giard) Fetch, var. major Fetch, 

‘ fresenii Nowak., 287 
grylli (Fres.) Nowak., 287 
muscae Cohn, 286 
papillata Thaxt, 287 
planchoniana (Cornu) Fetch, 287 
planchoniana Thaxt, 287 
tenthredinis (Fres.) Thaxt, 287 
thaxteriana Fetch, 287 
tipulae (Fres.) Fetch, 286 
Endomycopsis albicans (Vuili.) Dekker, 246 
Endothia Fr., 277 
Entoloma (Fr.) Quel, 205, 206 
bulbigenum B. & Br., 205, 206, 208 


Entoloma {continued) 
fertile Berk.j 192 

griseocyaneum (Fr.) QueL, 207, 208 
liquescens Cooke, 192 
lividum (Bull.) Fr., 350 
thomsoni B. & Br., 206, 208 
wynniae B. & Br., 206, 208 
de Bary, 346 

Entomophthora Fres., 298, 300, 301 
americana Thaxt., 289 
anglica Fetch, 289 
aphidis Hoffm., 288 
aphrophorae Rostr., 288 
(Tarichium) atrosperma Fetch, 290 
coleopterorum Fetch, 289 
dipterigena Thaxt., 289 
echinospora Thaxt., 289 
ferruginea Phillips, 287 
grylli Fres., 287 
lampyridarum Thaxt., 289 
lauxaniae Bubak, 290 
muscivora Schroet., 288 
occidentalis Thaxt., 288 
planchoniana Cornu, 287 
reticulata Fetch, 29a 
(Tarichium) richteri (Bres. & Staritz) Bubak, 

sphaerosperma Fres., 288, 300, 301 
tenthredinis Fres., 287 
tipulae Fres., 286 
variabilis Thaxt., 289 
virescens Thaxt., 290 
Entyloma de Bary, 346 
Epkhloe, the water relations of spore discharge 
in, by C. T. Ingold, 277 
Epichloe (Fr.) TuL, 277, 279 

typhina (Pers. ex Fr.) TuL, 277, 280 
Equisetum palustre, 143 
Eriophyes ribis, 293 
Erirrhinus nereis, 293 
Erysiphe cichoracearum DC., 143 
graminis DC., 54-56, 58-64, 349 
Escherischia coli (Migula) Castellani & 
Chalmers, 210 
Euastrum ansatum, 80 
Euchelia jacobaeae (cinnabar moth), 292 
Eutypa flavovirens (Fr.) TuL, 143 
Exhibit of classical British moulds, from tlie 
Herbarium of the Imperial Mycological 
Institute, 357 

Exobasidium vexans Massee, 229, 232, 233 
Exobasidium vexans, the basidiospores of, by 
C. H, Gadd & G. A. Loos, 229 

Favolaschia bispora, 36 
Festuca elatior, 249 
pratensis, 249 
Ficus carica (fig) , 86 

Filipendula ulmaria (meadow sweet), 143, 287 
Fimbriaria, 9 
angusta, 7 

Findlay, W. F. IC., * The use of perforated cards 
for preliminary identification of fungi 1 06 
Flammula dilepsis B. & Br., 25 
Forays, Bookham Common, 1946, 350 
Harrow Weald Common, 1946, 350 


London, 1946, 350 
Oxshott, 1946, 350 
Ruislip, 1946, 350 
Sandsend, 1946, 349 
Forficula (earwig), 287 
Fragaria (strawberry), 287, 293, 341 
Fraxinus (ash), 142, 292 
Fungi, genetics and cytology, discussion, 353 
Fusarium, Link ex Fr., 282, 346 
caeruleum (Lib.) Sacc., 247 
graminearum Schwabe, 247 
lateritium Nees, 87 
lini Bolley, 341 

oxysporum Fr., f. lini (Bolley) Snyder & 
Hansen, 341 

Fusicladium lini Sorauer, 90 

Gadd, C. H, & Loos, G. A., ‘The ambrosia 
fungus of Xyleborus fornicatus Eich.,’ 13 
‘ The basidiospores of Exobasidium vexans ’,229 
Galeruceiia tenelia, 294 
tenera, 290 
Galium aparine, 143 
Geaster fornicatus (Huds.) Fr., 350 
Gibberella baccata (Wailr.) Sacc., 87 
var. moricola (de Not.) Wollenw,, 87 
moricola (de Not.) Sacc., 86, 87 
zeae (Schw.) Fetch, 143 
Gibelluia aranearum (Schw.) Fetch, 297 
aranearum (Schw.) Syd., 143 
aranearum Syd., 297 
Gladiolus, 341 

Gloeosporium Desm. & Mont., 346 
perennans Zeller & Childs, 340 
Gloniopsis ievantica Rehm, 143 
* Glutinium macrosporum Zeller, 88, 89 
Giyceria, 142 
maxima, 242, 143, 290 
Gnomonia Ges. & de Not., 341 
Gonatosporium puccinioides (Fr.) Corda, 

. 143 

Gongylidium rufipes, 290, 297 
Grainger, J., ‘The ecology^ of Erysiphe graminis 

DG.% 54 

Gramineae (grasses), 341 
Gyrosigma, loi 

Harris, G. C. M,, ‘The morpholog)?- of 
Penkillium chrysogenum in submerged cul- 
ture’, 356 

. Helicosporium phragniitis v. Hoehn., 142, 

'Heiiomyces car^rotae Berk., 23 
leveillianus Berk., 28 

Helminthosporium Link ex Fr., 249, 251, 252, 


broroi Died., 249 
dictyoides DrecfasL, 249 
■gramineum.Rabenh.,' 136-138 
monoceras DrechsL, ■ 1 36-1 38 
sativum Pamm., King 8 c Bakke, 136, 137, 

.,139 ' . , 

siccans DreschL, 249, 252 
teres Sacc., 249 
Hepialus, 291, 294, 295, 

Hesperethusa crenulata, i 



Heterosporium Klotzsch ex Cooke, 346 
ailii EIL & Martin var. cepivorum Nicolas & 
. .Aggery, 341 

iridis (Faut. & Roum.) Jacques, 347 
monte^inum Bub., 347 
variabile Cooke, 341 
Hieracium, 116 
murorum, 1 1 6 
pilosella, 116 
Hirsutella Pat, 299 

acridiorum Fetch, 298, 304 
aphidis Fetch, 298 
citriformis Speare, 297, 304 
dipterigena Fetch, 298, 299 
eleutheratorum (Nees) Fetch, 291, 297, 304 
exoleta (Fr.) Fetch, 297 
lecaniicola (Jaap) Fetch, 292, 298 
saussurei (Cooke) Speare, 297, 303 
subulata Fetch, 297, 303 
Hoicus mollis, 277 
Hordeum (barley), 340 
Horse hair blight, 32, 34 
Hydnum (Linn.) Fr., 235 
ferruginosum Fr., 235 

Hygrophorus agathosmus (Fr. ex Secretan) 
Fr., 196, 208 
aromaticus Berk., 192 
cerasinus Berk., 196, 208 
foetens Phillips, 201, 208 
micaceus B. & Br., 196, 208 
obrussens Fr,, 20 
ventrkosiis B. & Br., 197, 208 
virgineus (Wuif.) Fr., 197, 208, 349 
Hymenochaete Lev., 235, 241, 244 
cinnamomeum (Pers.) Bres., 235 
Hymenostilbe Fetch, 304 
arachnophila (Ditm.) Fetch, 296 
formicarum Fetch, 304 
meianopoda (Speg.) Fetch, 304 
muscaria Fetch, 292, 296 
sphecophiia (Ditm.) Fetch, 291, 296 
sphingum (Schw.) Fetch, 296 
Hypomyces solani Reinke & Berth., f. cucur- 
bitae, 77 

Hypoxylon Bull, ex Fr., 277 
rubiginosum. (Pers.) Fr., 143 

Impatiens balsamina, 90 
Index, British agarics and^ boleti, 147 
Index, British fungi parasitic on lichens, 333 
Ingold, G. T., ‘The water-relations of spore 
discharge in Epichloe\ 277 
Inocybe fibrosa (Sow.) Fr., 205, 208 
International herbarium of plant pathogenic 
fungi, suggestions for, by P. Neergaard, 

. 3 # . 

Ins germamca, 87, 347 
pumila forrnosa, 87 
Isaria Pers. ex Fr., 295 , 
arachnophila Ditm., 296 
aranearum Schw., '297 
densa (Link) Fr., 302 
dufoia Delacr., 291, 295 
eleutheratorum Nees, 297 
exoleta IT., 297 

farinosa (Holmsk.) Fr,, 143, 295 

floccosa Fr., 296, 297 
fumoso-rosea (Wize) Vassil, 302 
guignardi Maheu, 304 
ochracea Bond, 296, 302, 303 
sphecophiia Ditm., 296 
sphingum Schw., 291, 296 
stilbxformis Speg., 304 
strigosa Fr., 296 

tenuipes Peck, 296, 299, 302, 303 

Jefferys, E. G., ‘A technique for rapid demon- 
stration of the production of antifungal 
substances by fungi or other micro- 
organisms’, 246 

Jubilee Volume; Editorial note, 144 
Juglans (walnut), 341 

Kelisia scottii, 297, 304 

La Touche, G. J., ‘Slide- traps for soil fungi’, 

Laccaria nana Massee, 196, 208 
Lachnocladium Lev., 235, 241, 244 
Lactarius Pers. ex S. F. Gray, 287, 350 
deliciosus (Linn.) Fr., 349 
quietus Fr., 35a 
subdulcis (Pers.) Fr., 350 
Lactuca (lettuce), 112, 124, 288 
muralis, 116 
saligna, 116 
scariola, 116 
Lagerstroemia, 37 

Lampyris noctiiuca (glow-worm), 295 
Laschia bispora (Holterm.) Sacc. & Trav., 36 
Lecanium, 292, 298 
Lentinus Fr., 19 
Leotia lubrica Fr., 350 
Lepidosaphes, 293 
ulmi, 295 

Lepiota amianthina (Scop.) Fr., 193, 208 
clypeolarioides Rea, 192, 208 
emplastrum Cooke & Massee, 192 
polysticta Berk., 192, 208 
Leptonia chalybea (Pers.) Fr., 201 
Leptosphaeria Ces. & de Not., 9, 1 1 
heterospora (de Not.) Niessl, 87 
*porellae Thirumalachar, n.sp., ii 
Leuciscus rutilus, 254, 266 
Lichens, list of British fungi parasitic on, by 
W. Watson, 305 

Linum usitatissimum (flax), 88, 90, 341 
Living mycelium, an improved technique for 
the staining of, by J. A. Macdonald, 92 
Lochmaea suturaiis, 289 
Lolium italicum, 249, 2 5 1 , 2 52 
multiflorum, 249 
perenne, 249 

Lophiostoma salicinum (Fabr.) Sacc., 143 
Lophodermium arundinaceum (Fr.) Ghev., 143 
Lycopersicum esculentum (tomato), 341 
Lygus pabuHnus, 286 

Macdonald, J. A., ‘An improved technique for 
the study of living mycelium’, 92 
McKay, R.j ‘Perithecial development of 
Vmturia inaequalis on scabbed apples 285 



Macrosiphon taraxaci, 287 
Marasmius Fr., 19, 38, 205 
actinophorus B. Br., 34 
*aibocapitatus Fetch, n.sp., 31, 41 
*alliarius Fetch, n.sp., 37, 43 
*amomi Fetch, n.sp., 37, 43 
androsaceus (L.) Fr., 92, 93 
"^ascendens Fetch, n.sp., 34, 42 
atrorubens Berk., 29 
*brunneostriatus Fetch, n.sp., 24, 40 
’^calami Fetch, n.sp., 38, 43 
caivus B. & Br., 20, 23 
campanella Holterm., 35, 36, 39 
»caryotas (Berk.) Fetch, 23 
chondripes B. &. Br., 29, 30 
*cineraceus Fetch, n.sp., 28, 41 
concolor, B. & C., 38 
confertus B. & Br., 29, 30 
congregatus Mont., 22, 23 
coniatus B. & Br,, 30, 31 
conigenus (Fers.) Karst., 199 
cornicoior B. &. Br., 21 
coronatus Fetch, 34 
corticigena B, & Br., 27 
crispatus B. &. Br., 23 
epochnous B. & Br., 37, 38 
equicrinus Muell., 33 
esculentus (VVulf.) Karst., 199 
eximius B. & Br., 26 
florideus B. & Br., 29 
fulviceps Berk., 24 
(Mycena) galericuia Ges., 39 
gordipes Sacc. & Paoi., 32 
♦griseoviolaceus Fetch, n.sp., 33, 42 
haematocephaius Mont., 24, 30 
*hakgaiensis Fetch, n.sp., 35, 42 
*var. denudata Fetch, n.var., 35, 43 
helvoius Berk. var. brnnneolus B. & Br., 30 
hemibaphus B. & Br., 29, 30 
hirtellus B. & Br,, 25 
hypociinoides B. & Br., 23 
ignobilis B. &. Br., 37 
inustus B. &. Br., 39 
*lateritius Fetch, n.sp., 31, 41 
leucophaeus (B. & Br.) Fetch, 21, 40 
ieveillianus (Berk.) Pat., 28 
menierie Bond,, 143 ' 

^micraster Fetch, n.sp., 32, 42 
multiju^s (B. &. Br.) Fetch, 20 
mntabilis B. & Br., 25, 26 
nephelodes (B. & Br.) Fetch, 19 
nivosus Berk., 27 
nummularins B. & Br., 24 
obscnratus B. Br., 25, 34 
obscuratus Fetch non B, & Br., 34 
ochracens B. & Br., 19 
paliidorubens (B. & Br.) Fetch, 24 
*paspali Fetch, n.sp., 35, 43 
pellucidus B. & Br., 22 
’**porphyreticus Fetch, n.sp., 36, 43 
prasinns, 39 
proletarins B, & G., 39 
proximus B. & Br., 25, 34 
pulcher (B. & Br.) Fetch, 38 
^pnrpureo-albus Fetch, n.sp., 37, 43 
*purpureo-grisens Fetch, n.sp., 20, 40 

radians B. & Br., 39 
ramealis (Bull.) Fr., 350 
*rigidichorda Fetch, n.sp., 34, 42 
runilosus B. & Br., 20, 21 
retails B. & Br., 33, 34 
rufescens B. & Br., 35 
***rufo-aurantiacus Fetch, n.sp., 36, 43 
*rufo-ochraceus Fetch, n.sp., 22, 40 
semipeliucidus B. & Br., 31 
**‘senescens Fetch, n.sp., 26, 41 
**‘stypinoides Fetch, n.sp., 28, 41 
stypinus B. & Br., 28 
subcinereus B. & Br,, 20, 25 
*subconiatus Fetch, n.sp., 31, 41 
suiciceps Berk., 40 
tiiwaitesii B. & 32 

*tubulatus Fetch, n.sp., 33, 42 
umbraculum B. & Br., 28 
■vv’^ainei B. & Br., var, auroricolor B. & Br., 

Marchantia, 7, 9 
Marssonina Magn., 346 
Massospora richteri Bres. <& Staritz, 290 
Matiier, K., ^Geneticai flexibility and the 
breeding system in fungi 355 
Melanospora parasitica TuL, 299 
Meliola os>Tidicola Hansf., 2 
Melolontha vulgaris (cockchafer), 294, 302 
Melosira, loi 

MetaiThizium anisopilae (Metsch.) Sorok, 294 
glutinosum S. Pope, 247 
Microcera coccophiia Desm., 298 
Microsphaera alni (DC.) Wint,, 63 
alphitoides GriH*. & MaubL, 341 
Mitruia scierotipus Boud., 143 
Monacrosporium Oudem., 14, 15, 17 

*ambrosium Gadd & Loos, n.sp., 16, 17, 18 
Monilia penicillioides Delacr., 292 
Moore, W. C., ‘British fungi’, 86 
‘Review: Flax diseases by R. McKay’, 144 
Moms, 87 
alba, 87 

Mougeotia, 94-96, 128, 130 
Mucor hiemaiis Wehiner, 354 
Mushroom, 126, 341, 356 
Mycena (Fr.) S. F. Gray, 40, 198, 200, 351, 352 
*auroricoior Fetch, n.nom., 40 
berkeleyi Massee, 191, 199, 208 
capiliaripes Feck, 352 
chlorantha Fr., 352 
consimiiis Cooke, 192 
fllopes (Bull.) Fr., 352 
gaiericulata (Scop.) Fr., 39, 199, 208 . 
iodioiens Lundeli, 352 
pallido-rubens, 25 
paupercula Berk., 199, 208 
rubromarginata Fr., 352 ' 
vi'tilis Fr., 352 ■ 

Mycogone Link ex Ghev., ^346 ' 
Mycosphaereila Johanson,. 346 
Myosotis, 143 

Myriangiunx duriaei Mont. & Berk., 292 . 
Myrica gale, 143, ■ 

Myrophagus ucrainicus (Wize) Sparrow, 290 
observations on, thC' genus*. II, by 
N. G*. Preston, 271 . 


Myrothecium Tode, 271 
gramineum Lib., 271, 272 
inundatum Fr., 271, 272 
^joUymannii N. C. Preston, n.sp., 272 
roridum Tode ex Fr., 272, 275, 341 
*striatosporum N. G. Preston, n.sp., 272, 275 
verrucaria (Alb. & Schw.) Ditm.,^75 
Myzocytium irregulare H. E. Petersen, 84 
megastomura De Wildem., 80^84 
proliferum Schenk, 84 

Nabis lineatus, 297, 304 
Nadsonia richteri Kostka, 76 
Napicladium arundinaceum (Gorda) Sacc., 143 
Naucoria Fr., 40 
furfuracea Pers., 25 
Navxcula, loi 
Nectria Fr., 277, 346 

Neergaard, P., ‘Suggestions for an Inter- 
national Herbarium of Plant Pathogenic 
Fungi’, 346 
Nerium, 3 
oleander, 3, 5 

Neurospora crassa Shear & Dodge, 356 
Newby, H. V., ‘Observations on Saprolegni- 
aceae. I. Saprokgtiia anisospora de Bary’, 

‘Observations on Saprolegniaceae. 11 . 
Saprolegnia paradoxa Maurizio’, 266 
Nicotiana tabacum (tobacco), 272, 274, 275 
Nitschkia cupularis (Pers. ex Fr.) Karst., 143 
Nitzschia sigmoidea, 10 1 
Nolanea babingtonii B. & Br., 206, 208 
llilvo-strigosa B. & Br., 192!Ia (Winter) Sacc., 206, 208 
rubida Berk., 192 
strigosissima Rea, 206 
Nomuraea.prasina MaubL, 295 
Noreilia upinimana, 296 
Notothylas, 9 

Nowakowskiella Schroet., 7 
ramosa Butler, 7 
Nudaria seiiex, 293 

Observations on Saprolegniaceae. I, by H. V. 
Newby, 254 

Observations on Saprolegniaceae. II, by 
H. V. New-by, 266 

Observations on the nutritional requirements 
Streptomyces griseus (Krainsky) Waksman 
& Schatz^ by J. F. Spilsbury, 210 
Odonestis potatoria, 292 
Oedogonium, 96 
Oidiuni link ex Fr., i, 86 
lini Skoric, 341 
Olpidiopsis Cornu, 9 
aphanomycis Cornu, 9 
ricciae Du Plessis, 9 
ucraiiiica Wize, 290 
Olpidium immersum Sorok., 84 
Omphalia (Fr.) S. F. Gray, 201, 352 
abhorrens B. & Br., 200, 208 
affricatus Fr., 201 , 
aluiacea Cooke & .Massee, 201, 208 
atropuncta (Pers.) Quci., 197, 208 
delicia B. & Br., 25 


directa B & Br., 192 
infumata B. & Br., 202, 208 
telmatiaea B. & C., 201, 208 
Oospora lactis (Fres.) Sacc., 354 
ovorum Trabut, 292, 302 
Ophiobolus porphyrogonus (Tode) Sacc., 88 
rubellus (Pers. ex Fr.) Sacc., 88 
Ophiocordyceps clavulata (Schw.) Fetch, 298 
Osyris wightiana, 2, 3 
Oudemansiella subaurantiaca, 37 
Ovularia Sacc., 346 

destructiva (Phill. & Plowr.) Massee, 143 

Pachybasium hamatum (Bon.) Sacc., 143 
Pachydiscia marchantiae (Berk.) Boud., 143 
Paederis riparius, 297 
Paspalum digitatum, 35 
Passalora graminis (Fuckel) v. Hoehn., 143 
Paxillus involutus (Batsch.) Fr., 350 
Pearson, A. A. & Dennis, R. W. G., ‘Revised 
list of British agarics and boleti’, 145 
Pemphigus lactucarius, 288 
Penicillium Link ex Fr., 30, 77 
chrysogenumThom, 211, 356 
expansum (Link) Thom, 136 
gladioli McCull. & Thom, 247 
janczewskii ZaL, 247 

notatum Westling, 66, 67, 74, 76, 21 1, 354 
patuluni Bainier, 136 

Penicillium notatum, spontaneous variation in, 
by E. R. Sansome, 66 
Peniophora Cooke, 143 
aegerita von Hoehn. & Litsch., 143 
Candida Lyman, 143 
leprosa Bourd. & Galz., 143 
pubera (Fr.) Sacc., 143 
Pennisetum subangustum, 271 
Pentatrichopus fragariae, 287 
Pergamasus crassipes, 287 
Peronospora Gorda, 346 
calotheca de Bary, 143 
dipsaci TuL, 86 
Pestalozzia de Not., 346 
Fetch, T., ‘A revision of Ceylon Marsamii’, 19 
‘A revised list of British entomogenous fungi 

Peucedanum palustre, 143 
Pezicula pruinosa Farlow, 89 
Peziza aurantia Pers., 350 
Phaeosphaerella ricciae F. L. Stephens, 7 
Phalaris arundinacea, 142 
Philaenus leucophthalmus, 287, 288 
spumarius, 288 
Phlyctidium Wallr., 94, 95 

*apophysatum H. M. Canter, n.sp., 94, 95 
Phlyctochytrium Schroet., 94, 98, 100 
Phoiiota erinacea (Fr.) Quel., 143 
PhomaDesm., 11,87,346 
Phomopsis viticola Sacc., 88 
Phragmites, 142 
communis, 143 

Phyllosticta Pers. ex Desm., 346 
camelHae Westend., 88 
camelliaecola Brun., 88 
var. meranensis Bub^k, 88 
dahliicola Brun., 90 


Phytophthora de Bary, 346 
cryptogea Pethybr. & Laff., 89 
infestans (Mont.) de Bary, 46 
Picromeris bidens, 294 
Pieris brassicae, 288, 300 
Piimularia, loi 
Pinus sylvestris, 235 
Pistillaria aculeata Pat., 143 
Pisum, 290, 301 
Piagiochasma, 9 
Plant pathology, disease measurement in, 

Plant Pathology Field Day, 1947, 351 
Plasmodiophora Woronin., 346 
Plasmopara nivea (Ung.) Schroet., 143 
Plateumaris braccata, 293, 294, 299 
Pleurage anserina (Ces.) Kuntze, 353 
Pleurotaenium Ehrenbergii, 80 
Pleurotus hobsoni Berk., 202, 208 
iaurocerasi B. & Br., 191, 203, 208 
palmatus (Bull.) Fr., 203 
petaloides (Bull.) Fr., 203, 208 
ruthae B. & Br., 203, 208 
tremens Quel., 203 
Plusia gamma, 290, 301 
Plutella macuiipennis, 300 
Pluteus Fr., 206, 208 
cinereus Quel., 206, 208 
spilopus B. & Br., 203, 205, 208 
*thomsoni (Berk.) Dennis, n.comb., 206, 208 
Poa trivialis, 143 

Podosphaera oxyacantliae (DC.) de Bar)% 56, 


PoHstes, 303 

Polyporus Mich, ex Fr., 346 
Pontecorvo, G., ‘Genetical aspects of hetero- 
karyosis’, 355 
Porella, 9, ii 

Poria Pers. ex S. F. Gray, i n 
Potato blight epidemics, the dependence on 
the weather of the date of outbreak of, by 
A. Beaumont, 45 

Potato blight, measurement of, 140 
President, C. G. G. Chesters, 357 
A. E, Miiskett, 357 

Preston, N. C., ‘Observations on the genus 
Myrothecium, II. Myrothecium grammum 
Lib., and two new species’, 271 
Primula juiiae, 90 
wanda, 90 
Proceedings, 353 
Prunus (cherry), 340 
Prunus (plum), 341 
padus, 63, 287 

Psalliota campestris (Linn.) Fr., 126 
Psathyrella disseminata (Pers.) Fr., 349 
Puccinia Pers., 353 
anomala Rostr., 340 
, graminis Pers., 353 
helianthi Schw., 353 
hordei Fuckel, 340 
hordei Otth, 340 
malvacearum Mont, 353 
persistens Plowr., 143 
simplex (Koern.) Eriks. & Hann., 340 
Pyrenopeziza arundinacea (DC.) Bond., 143 


Pyrenophoray a new species from Italian rye- 
grass, by H. F. Dovaston, 249 
Pyrenophora avenae Ito & 'Kuribay, 251, 252 
bromi Drechsl., 252 
*lolii Dovaston, n.sp., 251, 252 
teres Drechsl, 251, 252 
PyTonema confluens (Pers.) TuL, 353 
P>’Tus (apple), 340, 350 _ 

Pythiomorpha gonapodyides H. E. Petersen, 2 1 1 
Pythium Pringsh., 136, 137, 346 

aphanidermatum (Edson) Fitzp., 340 
de baryanum Hesse, 136 
ultimum Trow, 137, 340 
violae Chesters & Hickman, 341 

Quercus (oak), 63, 341 
robur, 341 

Ramuiaria Sacc., 346 
calthae (Erikss.) Lindr., 143 
Ranunculus repens, 288 
Receipts and payments, 358 
Review: Flax diseases by R. McKay, by 
W. C. Moore, 144 

Morth American Species of Mycena by A. H. 
Smith, by E. M. W^akefield, 351 
Rhizidium niycophiluin A- Braun, 98 
mycophilum Nowak., 98 
nowakowskii Karling, 98 
’^‘variabile H. M. Canter, n.sp., 98-100 
Rhizoctonia DC. ex Fr., 117, 281, 346 
soiani Kuebn, 136, 137, 246, 342 
Rhizophagus ferrugineus, 297 
Rhizophidium Schenk, 96 

^columnaris H. M. Canter, n.sp., 132-5 
globosum (A. Braun) Rabenhorst, 96, 97 
*variabiiis H. M. Canter, a.sp., 98 
Rhodes, A. F. Parker, ‘ Phenogenesis in fungi’, 
354 ^ 

Rhododendron, 341 
Rhodophyiius Quel., 206, 207, 351 
Rhoicosphenia cur\"ata, 102 
Rhytisma Fr., 346 
acerinum Fr., 143 
Ribes nigrum (black currant), 293 
Riccia, 9 

hinialayensis, 7 
Ridnus (castor oil plant), 13 
communis, 17, 18 
Robiiiia pseudacacia, 87 
Robinow, C., ‘On the staining of nuclei in 
bacteria and fungi 354 
Rosa (rose), 63 

Roseiliiiia ac|ui!a (Fr.) de Not., 143 
Russula Pers. ex S. F. Gray, 108, 350 
adusta (Pers.) Fr., 143 

Saccharomyces cerevisiae .Mcyen ex Hansen, 

. 354 ■ ■ ■ ■ ■ 

Salix (sallow, willow) , , 1 42 , 1 43 , 298 
SambuGus, 349 

Sansome,^E. R., ‘Spontaneous variation in 
PenkiUium notatum Strain N.R.R.L. 1249 
B21 66 , 

‘Mutation in Fungi imperfecti with special 
reference to FeniciUikm \ 336 



Santalum, 2 
album, 1,2 

Saproiegnia aoisospora de Bary, 254, 264, 269 
paradoxa Maurizio, 266, 267 
Saproiegniaceae, Observations on, by H. V. 

Newby, ,254, 266 
Sciara, 287 

Sclerotinia Fuckel, 346 
Sclerotium rolfsii Sacc., 136, 138 
Scolecotrichuni cladosporoideura Maire, 347 
iridis Faut. & Roum., 347 
Scopulariopsis peniciliioides (Delacr.) A. L. 

Smith & Ramsb., 292 
Senecio, 116 
jacobaea, 116 
vulgaris, 1 14, n6 
Septoria Sacc., 87, 346 
Serratia Bizio, 76 
marcescens Bizio, 75 
Sitones flavescens, 289 

Smith, G., h\nti-fungal substances from 
moulds. II. The effect of patulin on the 
growth of various plant parasites’, 136 
‘Autumn Forays, 1946’, 349 
Soil fungi, slide traps for, by C. J. La Touche, 

Solanum tuberosum (potato), 341 
Sonchus, 1 1 6 
oieraceus, 1 1 6 

Sorosporella uvella (Krass.) Giard, 298 
Sphacehma diseases of economic plants in 
Mysore, by M. J, Thirumalachar, i 
Sphaceioma de Bary, 1-3, 5, 6, 346 
’^curcumae Thirumalachar, n.sp., 6 
fawcettii Jenkins, i 
’^‘oleaiideri Thirumalachar, n.sp., 5 
*osyridis Thirumalachar, n.sp., 3 
’^'saiitali Thirumalachar, n.sp., 2 
Sphaeroderma fusisporum Fetch, 299 
pruinosum Peck, 88, 89 
Sphaeropsis malorum Peck, 76 
Sphaerotheca luliginea (Schlecht.) Saim., 86 
pannosa (VVallr.) Lev., 63 
Sphagnum, 201 
Sphinx iigustri, 296, 302 
Spicaria Harz, 295 

(Jsaria) farinosa (Holmsk.) VuilL, 291, 295, 

fumoso-rosea (Wize) V’assii.,,295 
prasina (Maubl.) Saw., 295 
stricta Fetch, 295 

(Isaria) swantoni (A. L. Smith) Fetch, 295 
Spilsbur)', J. F., ‘Observations on the nutri- 
tionai requirements of Streptomyces griseus 
(Krainsky) Waksman & Schatz’, 210 
Spinacia (spinach), 341 
Spirogyra, 98 

Spii'-otaeuia condensata, 80, 130, 135 
Sporo trichum aranearum Gav., 292 
isariae Fetch, 299 
Spring Foray, 1946, 142 
S tachybotiys atra Corda, 247 , 

Stagoiiospora Sacc., 89, 346 
Siainnaria persooni (Fr.) Fuckel, 143 
Staurastruin hinatum, 84 
Stciaphyiiuin W'aiir., 247, 346 

Stereum purpureum Pers., 247 
Stilbella Lindau, 299 
arridtii Lingelsh., 299 
kervillei (Quel.) Lingelsh., 299 
ramosa (Peck) Fetch, 299 
petiformis (Vahl) Fetch, 299 
Stilbum kervillei Quel., 299 
ramosum Peck, 299 

Streptomyces griseus (Krainsky) Waksman & 
Schatz, 210, 2 1 1, 216, 217, 219, 223, 225, 

Stromatinia (Bond.) Boud., 356 
Strophosomus lateralis, 294 
Studies on British Ghytrids, 11 , by H. M. 
Canter, 94 

Studies on British Ghytrids, III, by H. M. 
Canter, 128 

Synchytrium aurem Schroet., 143 
Synedra, loi 

Syngliocladium aranearum Fetch, 298 
cleoni (Wize) Fetch, 298, 299 

Tarichium Cohn, 301 
megaspermum Cohn, 290, 301 
richteri (Bres. & Staritz) Lakon, 290 
Technique for rapid demonstration of the 
production of antifungal substances by 
fungi or other micro-organisms, by E. G. 
Jefferys, 246 

Tetraploa aristata B. & Br., 143 
Thalictrum flavum, 143 
Thanmidium elegans Link, 247 
Thirumalachar, M. J., ‘Some new Sphaceioma 
diseases of economic plants in Mysore’, i 
‘ Some fungal diseases of bryophytes in 
Mysore 7 

Thomas, P. T., ‘Some aspects of the cytology 
of fungi’, 356 
Thread blight, 38 

Torrubia sphingum (Schw.) Tub, 291 
Torrubiella albolanata Fetch, 290, 292 
albotomentosa Fetch, 290 
aranicida Boud., 290 
Torymus cyanimus, 293 
Tragopogon, 116 
Treasurer’s Accounts, 358 
Trichoderma Pers. ex Fr., 282 
lignorum (Tode) Harz, 246 
viride Pers. ex Fr., 136, 137, 139, 247 
Tricholoma (Fr.) Quel., 197, 198 
aggregatum (Schaeff.) Quel., 195, 208 
argyraceum (Bull.) Fr., 195 
atrosquamosum Chev., 195 
circumtectum Cooke & Massee, 193, 195? 

crassifolium (Berk.) Ricksen, 197 
duracinum Cooke, 192 
georgii (Glus.) Fr., 198 
immundum Berk., 197, 198, 208 
myomyces Fr., 195, 208 , 

scalpturatum (Fr.) Quel., 195 
scalpfuratum Fr. sensu Bresadola, 208 
tenuiceps Cooke & Massee, 1 95, 208 
Trichophyton Malmsten, 77 
mentagrophytes, 77 
Triticum (wheat), 7, 342 


Ulex (gorse), 86, 293 
europaeus, 86 

Uimus montana (wych elm), 292, 298 
Urophora solstitialis, 293 
Urtica (nettle), 291 
Ustilago avenae (Pers.) Rostr., 341 
salvei B. & Br., 341 
striiformis (Westend.) Niessl, 341 

Ventxiria de Not., 346^ 

Venturia tnaequalis^ peritliecial development 
on scabbed apples, by R. McKay, 285 
Venturia inaequalis (Cooke) Wint., 285 
Vermicularia Fr., 346 
Verpatina Wbetzel & Drayton, 143 
VerticUium disease of mushrooms new to Great 
Britain, by F. C. Atkins, 126 
Verticillium Nees ex Walk., 126, 292, 346, 


albo-atrum Reinke & Berth., 136, 138 
eriophytis (Massee) Sacc. & Trott., 293 
malthousei Ware, 1 26, 341 
^menisporoides Fetch, n.sp., 295, 299 
psalliotas Treschow, 126 
Vicia (bean), 340 
Viola, 341 
comuta, 272 


Visit: Brewery Research Department, Barclay 
Perkins & Co., 356 
Volutella melaloma B.&.Br., 143 

Wakefield, E, M. Review. Morih American 
Species of Mycena (A. H. Smith), 351 
Waliichia, 22 

Watson, W-, ‘ List of British fungi parasitic on 
lichens or wliich have been included as 
lichens (or vice versa), with some notes 
on their characters and distribution 305 
Whitehouse, H. L. K., ‘Evoluiionary aspects of 
heterothaliism and heterokatyosis’, 355 
Wild, H., ‘Downy mildew disease of the 
cultivated lettuce’, 1 12 

Xerotus tener B. & Br., 35 
Xyleborus fornicatior, 1 3 
Xyleborus fornicatus Eich., the ambrosia fungus 
of, by C. H. Gadd & C. A. Loos, 13 
Xyleborus fornicatus, 13, 14, 17, 18 

Zygnema, 130 

Zygorhizidium Loewenthal, 128, 132 
verrucosum Geitier, 1 28 
wiilei Loewenthal, 128, 130 
Zythia fragariae Laibach, 341